Quercy Phosphorites Formation

Quercy Phosphorites Formation
Quercy Phosphorites Formation
Stratigraphic range: latest Bartonian-Late Oligocene; ~38–25 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Type	Formation
Overlies	Fissure fillings in karstified Jurassic and Triassic rocks
Lithology
Primary	Phosphorite
Location
Coordinates	44°18′N 1°36′E / 44.3°N 1.6°E
Approximate paleocoordinates	42°48′N 2°00′W / 42.8°N 2.0°W
Region	Occitanie
Country	France
Type section
Named for	Quercy
Named by	Thévenin
Year defined	1903
	Quercy Phosphorites Formation (France)
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The Quercy Phosphorites Formation (French: Phosphorites du Quercy; Occitan: Fosforits de Quercy) is a geologic formation and lagerstätte in Occitanie, southern France. It preserves fossils dated to the Paleogene period (latest Bartonian to Late Oligocene),^[1]^[2] or MP16 to MP28 zones of the European land mammal age classification, ranging from approximately 38 to 25 Ma.

It qualifies as a Lagerstätte because beside a large variety of mammals, birds, turtles, crocodiles, flora and insects, it also preserves the soft tissues of amphibians and squamates, in addition to their articulated skeleton in what has been called natural mummies.^[3]

The genera Quercylurus, Quercymegapodius, Quercypsitta, Quercypodargus, Quercycerta and Quercygama, and species Mosaicomeryx quercyi, Robiacina quercyi, Palaeophyllophora quercyi, Archaeomys quercyi, Eomys quercyi, Eucricetodon quercyi and Tarnomys quercynus, as well as the lizards Paraplacosauriops quercyi and Pseudolacerta quercyini and the insect Palaeortona quercyensis were named after the formation.

Description

The first phosphate deposits in Quercy were discovered in 1869 and published by Daubré and Trutat independently in 1871.^[4] The first fossils from the formation were described by Delfortie (1872) and Gervais in the same year and extensively studied by Filhol from 1877 onwards. The first geologic investigation of the formation was performed by Thévenin in 1903, and apart from a description by Gèze in 1938, the paleontological richness was not studied until a team of researchers of the Universities of Montpellier and Paris visited the site in 1965.^[5]

The karstified phosphate deposits are found from the Lot and Célé river valleys in the north to the left bank of the Aveyron in the south and from the Villefranche Fault in the east to the lacustrine deposits of the Aquitaine Basin in the west. The formation is found in fissures (karst) incising Jurassic and Triassic rocks east of Cahors.^[6] The age of the fossiliferous unit, in which almost 12,000 specimens were found ranges from the MP16 to MP28 zones of the European land mammal age classification.^[5] These ages correspond to the latest Bartonian to Chattian, from about 38 to 25 Ma.^[7]

Paleontological significance

The Quercy Phosphorites Formation is a highly fossiliferous unit designated as a Lagerstätte due to the excellent preservation of fossils. The phosphorite conserves up to the nerves, digestive tract and stomach content,^[8] insect larvae and other elements of the paleobiology in the formation. Nearly all Quercy fly pupae were preserved as isolated endocasts, of which many were still covered by the puparium, the hardened skin of the last larval instar.^[9] The formation also straddles the Grande Coupure and shows diversity changes (number of species) of frog, salamander, lizard and snake fossil records across the formation.^[10] It is assumed that the Quercy arthropods fossilized by a rapid fixation by phosphate-rich water followed by encrustation and mineralization.^[11]

Fossil content

The following fossils have been reported from the formation:^[1]^[2]^[12]^[13]

Mammals

Apatemyidae

Genus	Species	Note
Chardinyus	C. sp.^[12]	An apatotheria.
Heterohyus	H. (Chardinyus) nanus^[12]	An apatemyid.
Heterohyus	H. (Gervaisyus) pygmaeus^[12]	An apatemyid.
Gervaisyus	G. sp.^[12]

Artiodactyls

Genus	Species	Location	Time	Material	Notes	Images
Bachitherium	B. guirounetensis^[13]
Bachitherium	B. lavocati^[13]
Cryptomeryx	C. gaudryi^[14]
Dichobune	D. sigei^[13]
Dichodon	D. vidalenci^[13]
Iberomeryx	I. matsoui^[13]
Mosaicomeryx	M. quercyi^[15]
Paroxacron	P. bergeri^[16]
Plesiomeryx	P. cadurcensis^[16]
Prodremotherium	P. elongatum^[15]
Pseudamphimeryx	P. salesmei^[13]
Robiacina	R. lavergnensis^[13]
Robiacina	R. quercyi^[13]
Tapirulus	T. perrierensis^[13]

Carnivora

Genus	Species	Location	Time	Material	Notes
Amphicynodon	A. typicus^[17]	Pech Crabet locality	Oligocene	Two hemi-mandibles	An early amphicynodontidae caniform
Cephalogale	C. sp.^[18]			A piece of hemi-mandible
Cynodyctis	C. lacustris neboulensis^[13]	Sainte Néboule		A mandible	An amphicynodontidae caniform.
Dinailurictis	D. bonali^[19]^[20]	La Tuque	Oligocene.	Several holotype consists of a fragment of the left upper canine, lower canine, distal right humerus, distal left ulna, and a proximal piece of right femur.	A nimravid felidae
Eofelis	E. edwardsii^[21]^[22]				A nimravid
	E. giganteus^[23]	Phosphorites du Quercy		A mandible
	cf. E. sp.^[24]
Eusmilus	E. bidentatus^[21]^[22]		Eocene/Oligocene	A edentulous hemi-mandible and a fragment of upper canine	A nimravid
Huntictis	H. minima	Gardiol 3 locality	Early Oligocene	A holotype consists of a dentition.	A small musteloid
Mustelictis	M. cf. major^[25]			A part of right hemi-mandible with alveolus of p1, p2, fragments of p3 roots, and p4-m1.
Mustelictis	M. aff. olivieri^[25]			A holotype specimen consists of a skull and a hemi-mandible.
Nimravus	N. intermedius^[19]^[20]	MP 22 localities, La Plante2, Mas de Got and Valbro localities	Oligocene	A left hemi-mandible with m1	A nimravid felidae.
Pachycynodon	P. amphictina^[26]		Oligocene	Multiple specimens consist of hemi-mandible and dentitions.
	P. crassirostris^[27]
	P. cf. dubius^[26]
	P. cf. filholi^[28]
Palaeogale	P. sectoria^[22]	Valbro locality	Oligocene	A specimen consists of p4 dentition.	An early feliformia
Peignictis	P. pseudamphictis^[29]	Valbro locality	Early Oligocene	A posterior part of hemi-mandible	A carnivoran with uncertain affinities
Quercylurus	Q. major^[19]	Phosphorites du Quercy	Oligocene	A maxilla	A nimravid
Storchictis	S. miacinus	Phosphorites du Quercy	Late Eocene	A mandible	An Amphicyonidae
Wangictis	W. tedfordi^[30]	Phosphorites du Quercy	Early Oligocene	A left hemi-mandible with c-m2	An Amphicynodontidae

Chiroptera

Genus	Species	Notes
Hipposideros	H. pseudorhinolophus sehlosseri^[12]	An early roundleaf bat.
	H. pseudorhinolophus trassounius^[12]
	H. pseudorhinolophus zbrjdi^[12]
Leuconoe	L. lavocati^[12]	An early vespertilionid bat.
Palaeophyllophora	P. oltina^[12]	An early hipposiderid bat.
Palaeophyllophora	P. quercyi^[12]	An early hipposiderid bat.
Stehlinia	S. bonisi^[12]	An early natalidae bat
Stehlinia	S. minor^[12]	An early natalidae bat
Vaylatsia	V. garouillasensis^[12]
Vespertiliavus	V. gerscheli^[12]	An early Emballonurid bat
	V. gracilis^[12]
	V. lapradensis^[12]

Creodonta

Genus	Species	Location	Time	Material	Images
Paracynohyaenodon	P. magnus^[12]
Parapterodon	P. lostangensis^[12]

Euarchonta

Genus	Species	Location	Time	Material	Images
Darbonetus	D. aubrelongensis^[31]

Hyaenodonta

Genus	Species	Location	Time	Material	Notes	Images
Paroxyaena	P. pavlovi^[32]	Phosphorites du Quercy	Late Eocene	A partial skull	A hyaenodont

Lipotyphla

Genus	Species	Location	Time	Material	Notes	Images
Amphidozotherium	A. cayluxi^[12]
Darbonetus	D. aubrelongensis^[12]
Darbonetus	D. tuberi^[12]
Myxomygale	M. antiqua^[33]
Saturninia	S. beata^[12]
Saturninia	S. pelissiei^[12]

Marsupials

Genus	Species	Location	Time
Amphiperatherium	A. bourdellense^[12]	Le Bretou Locality	Oligocene
	A. lamandini^[12]
	A. minutum
Peratherium	P. bretouense^[12]
	P. cayluxi^[12]
	P. lavergnense^[12]
	P. perrierense^[12]

Perissodactyls

Genus	Species	Location	Time	Material	Images
Pachynolophus	P. bretovense^[13]

Primates

Genus	Species	Location	Time	Material	Images
Cryptadapis	C. tertius^[12]			An adapiform primate.

Proeutheria

Genus	Species	Location	Time	Material	Notes	Images
Pseudorhyncocyon	P. cayluxi^[12]

Rodents

Genus	Species	Location	Time	Material	Notes	Images
Archaeomys	A. intermedius^[13]	Pech du Fraysse	Oligocene		A theridomyidae rodent
Archaeomys	A. quercyi^[13]		Oligocene		A theridomyidae rodent
Bernardia	B. marandati^[13]
Blainvillimys	B. gemellus^[13]
	B. gousnatensis^[13]
	B. langei^[13]^[34]
	B. rotundidens^[35]
Elfomys	E. medius^[13]
Eomys	E. gigas^[13]	Pech du Fraysse Locality	Late Oligocene	A specimen	An eomyidae rodent
	E. minus^[13]	Pech du Fraysse Locality		A specimen
	E. quercyi^[13]	Pech du Fraysse Locality		A specimen		E. quercyi
Eucricetodon	E. atavus^[34]
Eucricetodon	E. quercyi^[13]
Gliravus	G. garouillensis^[13]
Gliravus	G. itardiensis^[13]
Issiodoromys	I. limognensis^[13]
Issiodoromys	I. pauffiensis^[13]^[34]
Palaeosciurus	P. goti^[13]^[34]
Paradelomys	Paradelomys spelaeus^[13]
Patriotheridcmys	P. altus^[13]
	P. altus neboulensis^[13]
	P. sudrei^[13]
Pseudoltinomys	P. gaillardi^[34]
	P. major^[13]
	P. phosphoricus^[13]
Sciuromys	S. cayluxi^[36]
Sciuromys	S. rigali^[13]
Tarnomys	T. quercynus^[13]
Theridomys	T. ludensis^[13]

Birds

Genus	Species	Location	Time	Material	Note
Ameghinornis	A. minor^[12]	Phosphorites du Quercy	Late Eocene	A specimen	An early european seriema
Euronyctibius	E. kurochkini^[37]	Phosphorites du Quercy	Eocene	A right humerus	An early nightjar
Horusornis	H. vianeyliaudae^[12]	La Bouffie Locality	Eocene	A specimen	An early accipitriformes
Dynamopterus	D. velox^[12]	Near Caylus	Oligocene	Almost complete right humerus	An extinct cuckoo bird, synonymous with Dynamopterus velox
Itardiornis	I. hessae^[12]		Eocene/Oligocene	A specimen	A Messelornithidae bird
Leptoganga	L. sp.^[12]
Necrobyas	N. minimus^[12]
Nocturnavis	N. sp.^[12]
Palaeoglaux	P. perrierensis^[12]				An early owl
Palaeotodus	P. escampsiensis^[12]		Late Eocene	A specimen	An early todies
Palaeotodus	P. itardiensis^[12]		Early Oligocene	A specimen	An early todies
Paleseyvus	P. escampensis^[12]
Primocolius	P. sigei^[12]	Le Bretou locality	Eocene	A limb element consists of a practically complete left humerus
Primocolius	P. minor^[12]		Eocene	A near-complete right tarsometatarsal
Quercymegapodius	Q. brodkorbi^[12]				A Quercymegapodiidae bird
Quercypodargus	Q. olsoni^[37]
Quercypsitta	Q. ivani^[12]	La Bouffie locality	Late Eocene	A holotype specimen consists of three coracoids	An early offshoots of parrot
Quercypsitta	Q. sudrei^[12]	La Bouffie locality	Late Eocene	Fragmentary sets of postcrania, which consist of foot and wing bones	An early offshoots of parrot
Recurvirostra	R. sanctaeneboulae^[12]
Strigogyps	S. dubius	Phosphorites du Quercy	Late Eocene	A specimen
Sylphornis	S. bretouensis^[12]
Ventivorus	V. ragei^[12]	Le Bertou Locality			An early nightjar

Reptiles

Crocodilians

Genus	Species	Location	Time	Material	Notes	Images
Alligator	A. gaudryi^[38]

Lizards

Genus	Species	Location	Time	Material	Notes
Ayalasaurus	A. tenuis^[39]
Bauersaurus	B. cosensis^[40]	Cos Locality	Uppermost Early Eocene	The upper maxilla) and lower jaw fragments.	A pan-gekkotan lizard.
Brevisaurus	B. smithi^[39]
Cadurciguana	C. hoffstetteri^[12]
Cadurcogekko	C. piveteaui^[39]
Cadurcopanoplos	C. vaylatsensis^[41]	La Bouffie locality	Late Eocene	Two holotypes consist of a single incomplete frontal bone and a partial right mandible.	A Glyptosaurinae anguid.
Dracaenosaurus	D. croizeti^[39]
Eurheloderma	E. gallicum^[39]
Geiseltaliellus	G. lamandini^[39]
Mediolacerta	M. roceki^[39]
Necrosaurus	N. cayluxi^[39]
Necrosaurus	N. eucarinatus^[39]
Omoiotyphlops	O. priscus^[39]
Paraplacosauriops	P. quercyi^[39]
Phosphoriguana	P. peritechne^[41]	La Bouffie Locality	Late Eocene	A single partial lower jaw	A probable member of pleurodonta
Placosaurus	P. rugosus^[39]
Placosaurus	P. sp.^[39]
Plesiolacerta	P. lydekkeri^[39]
Pseudeumeces	P. cadurcensis^[39]
Pseudolacerta	P. mucronata^[39]
Pseudolacerta	P. quercyini^[39]
Quercycerta	Q. maxima^[39]
Quercygama	Q. galliae^[39]
Uromastyx	U. europaeus^[39]

Snakes

Genus	Species	Location	Time	Material	Notes	Images
Cadurcoboa	C. insolita^[12]
Coluber	C. cadurci^[12]
Dunnophis	D. cadurcensis^[12]
Eoanilius	E. europae^[12]
Natrix	N. mlynarskii^[12]^[42]
Palaeopython	P. cadurcensis^[38]
	P. filholi^[38]
	P. neglectus^[38]
Paleryx	P. cayluxi^[38]
Platyspondylia	P. lepta^[12]
Plesiotortrix	P. edwardsi^[38]

Turtles

Genus	Species	Location	Time	Material	Notes	Images
Ptychogaster	P. cayluxensis^[43]
Testudinidae	indeterminate ^[43]
Testudo	T. phosphoritarum^[38]

Amphibians

Frogs

Genus	Species	Location	Time	Material	Notes
Bufo	B. serratus^[38]	Phosphorites du Quercy	Eocene	A specimen	An early true toad.
Thaumastosaurus	T. bottii^[38]	Phosphorites du Quercy	Eocene	A specimen	An early Pyxicephalinae frog
Thaumastosaurus	T. gezei^[3]	Phosphorites du Quercy	Eocene	A mummified preserved skeleton	An early Pyxicephalinae frog

Salamanders

Genus	Species	Location	Time	Material	Notes	Images
Heteroclitotriton	H. zitteli^[38]	Phosphorites du Quercy locality	Late Eocene	A specimen	An early salamander
Megalotriton	M. filholi^[38]	Phosphorites du Quercy locality	Late Eocene	A specimen	An extinct salamander
Megalotriton	M. portisi^[38]	Phosphorites du Quercy locality	Late Eocene	A specimen	An extinct salamander
Phosphotriton	P. sigei^[44]	Phosphorites du Quercy locality	Late Eocene	A mummified skeleton.	A frog-eating salamander.	Exceptional preservation of nerves, digestive tract and stomachal content in the frog-eating salamander Phosphotriton sigei

Insects

Blattodea

Family	Genus	Species	Location	Age	Notes	Images
					Various cockroaches^[45]

Diptera

Family	Genus	Species	Location	Age	Notes	Images
					Various flies^[45]

Coleoptera

Family	Genus	Species	Location	Age	Notes	Images
Histeridae	†Onthophilus	†O. intermedius^[45]			A clown beetle
					Various beetles^[45]

Hymenoptera

Family	Genus	Species	Location	Age	Notes
Diapriidae	†Coptera	†C. anka^[46]	Near Bach	Paleogene	An early Diapriinae wasp
	†Palaeortona	†P. quercyensis^[46]	Near Bach	Paleogene	An early Diapriinae wasp
	†Xenomorphia	†X. handschini^[46]	Near Bach	Paleogene	An early Diapriinae wasp
	†Xenomorphia	†X. resurrecta^[9]	Near Bach	Paleogene	An early Diapriinae wasp
					Various wasps/ants/bees^[45]

Lepidoptera

Family	Genus	Species	Location	Age	Notes	Images
					Various butterflies/moths^[45]

Orthoptera

Family	Genus	Species	Location	Age	Notes	Images
					Various ensiferan grasshoppers/katydids^[45]

Myriapoda

Family	Genus	Species	Location	Age	Notes	Images
					Various centipedes and millipedes^[45]

Flora

Family	Genus	Species	Notes
Anacardiaceae	Indeterminate^[47]	Indeterminate^[47]	Indeterinate anacardiaceous fossils
Arecaceae	Palmocaulon^[48]	Palmocaulon sp.^[48]	A palm tree relative
Malvaceae	Indeterminate^[49]	Indeterminate^[49]	indeterminate malvaceous fossils
Menispermaceae	Indeterminate^[49]	Indeterminate^[49]	Indeterminate menispermaceous fossils
Sapotaceae	Indeterminate^[49]	Indeterminate^[49]	Indeterminate sapotaceous fossils
Ulmaceae	Indeterminate^[49]	Indeterminate^[49]	Indeterminate ulmaceous fossils
Vitaceae	Indeterminate^[47]	Indeterminate^[47]	Indeterminate vitaceous fossils
Indeterminate	Indeterminate^[49]	Indeterminate^[49]	Indeterminate caryophyllalean fossils

References

^ ^a ^b Quercy Phosphorites Formation at Fossilworks.org
^ ^a ^b Phosphorites du Quercy Formation at Fossilworks.org
^ ^a ^b Laloy et al., 2013
^ Legendre et al., 1997, p.331
^ ^a ^b Legendre et al., 1997, p.332
^ Astruc & Pellissié, 1988, p.8
^ Legendre et al., 1997, p.333
^ Tissier et al., 2017, p.6
^ ^a ^b Van de Kamp et al., 2018, p.2
^ Vasilyan, 2018, p.19
^ Van de Kamp et al., 2018, p.12
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah ^ai ^aj ^ak ^al ^am ^an ^ao ^ap ^aq ^ar ^as ^at ^au ^av ^aw ^ax ^ay ^az ^ba ^bb ^bc ^bd ^be ^bf Legendre et al., 1997, p.334
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah ^ai ^aj ^ak Legendre et al., 1997, p.335
^ Sudre, 1984
^ ^a ^b Mennecart & Métais, 2015
^ ^a ^b Blondel, 2005
^ Bonis et al., 2019, p.603
^ Bonis et al., 2019, p.612
^ ^a ^b ^c Peigné, 2003
^ ^a ^b Bonis et al., 2019, p.616
^ ^a ^b Peigné & Brunet, 2001
^ ^a ^b ^c Bonis et al., 2019, p.618
^ Peigné, 2000
^ Peigné, 2001
^ ^a ^b Bonis et al., 2019, p.614
^ ^a ^b Bonis et al., 2019, p.610
^ Bonis et al., 2019, p.604
^ Bonis et al., 2019, p.608
^ Bonis et al., 2019, p.615
^ Bonis et al., 2019, p.611
^ Hooker, 2018, p.236
^ Lavrov, 2007
^ Hugueney & Maridet, 2017
^ ^a ^b ^c ^d ^e Ginot et al., 2016, p.7
^ Hartenberger & Vianey-Liaud, 1978
^ Vianey-Liaud & Schmid, 2009
^ ^a ^b Mourer-Chauviré, 1989
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l Rage, 2006
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t Augé, 2005
^ Čerňanský, A.; Daza, J. D.; Tabuce, R.; Saxton, E.; Vidalenc, D. (2023). "An early Eocene pan-gekkotan from France could represent an extra squamate group that survived the K/Pg extinction". Acta Palaeontologica Polonica. 68 (4): 695–708. doi:10.4202/app.01083.2023.
^ ^a ^b Lemierre, Alfred; Georgalis, Georgios L. (2025). "Diversity in a greenhouse world: Herpetofauna from the late Eocene (MP 17A) of la Bouffie, Quercy Phosphorites (Lot, SW France)". Swiss Journal of Palaeontology. 144 40. doi:10.1186/s13358-025-00370-9.
^ Rage, 1988
^ ^a ^b Broin, 1977
^ Tissier et al., 2016
^ ^a ^b ^c ^d ^e ^f ^g ^h Schmied et al., 2013, p.145
^ ^a ^b ^c Van de Kamp et al., 2018, p.4
^ ^a ^b ^c ^d De Franceschi et al., 2006, p.100
^ ^a ^b De Franceschi et al., 2006, p.99
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j De Franceschi et al., 2006, p.101

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External links

Media related to Quercy Phosphorites Formation at Wikimedia Commons

[FWQuercy1-1] Quercy Phosphorites Formation at Fossilworks.org

[FWQuercy2-2] Phosphorites du Quercy Formation at Fossilworks.org

[Laloy2013-3] Laloy et al., 2013

[Legendre1997-p331-4] Legendre et al., 1997, p.331

[Legendre1997-p332-5] Legendre et al., 1997, p.332

[Astruc1988-p8-6] Astruc & Pellissié, 1988, p.8

[Legendre1997-p333-7] Legendre et al., 1997, p.333

[Tissier2017-p6-8] Tissier et al., 2017, p.6

[VanDeKamp2018-p2-9] Van de Kamp et al., 2018, p.2

[Vasilyan2018-p19-10] Vasilyan, 2018, p.19

[VanDeKamp2008-p12-11] Van de Kamp et al., 2018, p.12

[Legendre1997-p334-12] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah ^ai ^aj ^ak ^al ^am ^an ^ao ^ap ^aq ^ar ^as ^at ^au ^av ^aw ^ax ^ay ^az ^ba ^bb ^bc ^bd ^be ^bf Legendre et al., 1997, p.334

[Legendre1997-p335-13] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah ^ai ^aj ^ak Legendre et al., 1997, p.335

[Sudre1984-14] Sudre, 1984

[Mennecart2015-15] Mennecart & Métais, 2015

[Blondel2005-16] Blondel, 2005

[Bonis2019-p603-17] Bonis et al., 2019, p.603

[Bonis2019-p612-18] Bonis et al., 2019, p.612

[Peigne2003-19] Peigné, 2003

[Bonis2019-p616-20] Bonis et al., 2019, p.616

[PeigneBrunet2001-21] Peigné & Brunet, 2001

[Bonis2019-p618-22] Bonis et al., 2019, p.618

[Peigne2000-23] Peigné, 2000

[Peigne2001-24] Peigné, 2001

[Bonis2019-p614-25] Bonis et al., 2019, p.614

[Bonis2019-p610-26] Bonis et al., 2019, p.610

[Bonis2019-p604-27] Bonis et al., 2019, p.604

[Bonis2019-p608-28] Bonis et al., 2019, p.608

[Bonis2019-p615-29] Bonis et al., 2019, p.615

[Bonis2019-p611-30] Bonis et al., 2019, p.611

[Hooker2018-p236-31] Hooker, 2018, p.236

[Lavrov2007-32] Lavrov, 2007

[Hugueney2017-33] Hugueney & Maridet, 2017

[Ginot2016-p7-34] Ginot et al., 2016, p.7

[Hartenberger1978-35] Hartenberger & Vianey-Liaud, 1978

[Vianey2009-36] Vianey-Liaud & Schmid, 2009

[Mourer1989-37] Mourer-Chauviré, 1989

[Rage2006-38] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l Rage, 2006

[Auge2005-39] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t Augé, 2005

[40] Čerňanský, A.; Daza, J. D.; Tabuce, R.; Saxton, E.; Vidalenc, D. (2023). "An early Eocene pan-gekkotan from France could represent an extra squamate group that survived the K/Pg extinction". Acta Palaeontologica Polonica. 68 (4): 695–708. doi:10.4202/app.01083.2023.

[Lemierre2025-41] Lemierre, Alfred; Georgalis, Georgios L. (2025). "Diversity in a greenhouse world: Herpetofauna from the late Eocene (MP 17A) of la Bouffie, Quercy Phosphorites (Lot, SW France)". Swiss Journal of Palaeontology. 144 40. doi:10.1186/s13358-025-00370-9.

[Rage1988-42] Rage, 1988

[Broin1977-43] Broin, 1977

[Tissier2016-44] Tissier et al., 2016

[Schmied2013-p145-45] ^ ^a ^b ^c ^d ^e ^f ^g ^h Schmied et al., 2013, p.145

[VanDeKamp2018-p4-46] Van de Kamp et al., 2018, p.4

[DeFranceschi2006-p100-47] De Franceschi et al., 2006, p.100

[DeFranceschi2006-p99-48] De Franceschi et al., 2006, p.99

[DeFranceschi2006-p101-49] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j De Franceschi et al., 2006, p.101

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Quercy Phosphorites Formation

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