Lagerstätte

A Fossil-Lagerstätte (German: [ˈlaːɡɐˌʃtɛtə], from Lager 'storage, lair' Stätte 'place'; plural Lagerstätten) is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These formations may have resulted from carcass burial in an anoxic environment with minimal bacteria, thus delaying the decomposition of both gross and fine biological features until long after a durable impression was created in the surrounding matrix. Fossil-Lagerstätten span geological time from the Neoproterozoic era to the present.

Worldwide, some of the best examples of near-perfect fossilization are the Cambrian Maotianshan shales and Burgess Shale, the Ordovician Soom Shale, the Silurian Waukesha Biota, the Devonian Hunsrück Slates and Gogo Formation, the Carboniferous Mazon Creek, the Triassic Madygen Formation, the Jurassic Posidonia Shale and Solnhofen Limestone, the Cretaceous Yixian, Santana, & Agua Nueva formations and the Tanis Fossil Site, the Eocene Fur Formation, Green River Formation, Messel Formation & Monte Bolca, the Miocene Foulden Maar and Ashfall Fossil Beds, the Pliocene Gray Fossil Site, and the Pleistocene Naracoorte Caves & La Brea Tar Pits.

Types

Palaeontologists distinguish two kinds:^[1]^[2]

Konzentrat-Lagerstätten (concentration Lagerstätten) are deposits with a particular "concentration" of disarticulated organic hard parts, such as a bone bed. These Lagerstätten are less spectacular than the more famous Konservat-Lagerstätten. Their contents invariably display a large degree of time averaging, as the accumulation of bones in the absence of other sediment takes some time. Deposits with a high concentration of fossils that represent an in situ community, such as reefs or oyster beds, are not considered Lagerstätten.
Konservat-Lagerstätten (conservation Lagerstätten) are deposits known for the exceptional preservation of fossilized organisms or traces. The individual taphonomy of the fossils varies with the sites. Conservation Lagerstätten are crucial in elucidating important moments in the history and evolution of life. For example, the Burgess Shale of British Columbia is associated with the Cambrian explosion, and the Solnhofen limestone with the earliest known bird, Archaeopteryx.

Archaeopteryx specimen from Solnhofen Lagerstätte (Upper Jurassic)

Fossil specimen of Echinochimaera from the Bear Gulch Limestone (Middle Carboniferous)

Preservation

Konservat-Lagerstätten preserve lightly sclerotized and soft-bodied organisms or traces of organisms that are not otherwise preserved in the usual shelly and bony fossil record; thus, they offer more complete records of ancient biodiversity and behavior and enable some reconstruction of the palaeoecology of ancient aquatic communities. In 1986, Simon Conway Morris calculated only about 14% of genera in the Burgess Shale had possessed biomineralized tissues in life. The affinities of the shelly elements of conodonts were mysterious until the associated soft tissues were discovered near Edinburgh, Scotland, in the Granton Lower Oil Shale of the Carboniferous.^[3] Information from the broader range of organisms found in Lagerstätten have contributed to recent phylogenetic reconstructions of some major metazoan groups. Lagerstätten seem to be temporally autocorrelated, perhaps because global environmental factors such as climate might affect their deposition.^[4]

A number of taphonomic pathways may produce Konservat-Lagerstätten:^[5]

Phosphatization (replacing soft tissues with phosphate, such as Orsten-type and Doushantuo-type preservations).
Silicification (replacing or entombing soft tissues with silica, such as petrified wood or Bitter Springs-type preservation).
Kerogenization (soft tissues converted into inert carbonaceous films, as found in Burgess Shale-type preservation).
Aluminosilification (replacing or coating soft tissues with films of aluminosilicate minerals).
Pyritization (replacing soft tissues with pyrite, such as the exquisite detail found in Beecher's trilobite-type preservation).
Calcification (replacing or soft tissues with calcite minerals).
Siderite or calcite nodules (chemicals released by decaying soft tissue modify the surrounding sediment into a siderite concretion or coal ball encasing the fossil, as found in the Mazon Creek fossil beds).
Rapid sediment cementation (impressions of soft tissue preserved through casts and molds in the surrounding sediment, such as Ediacaran-type preservation facilitated by microbial mats).
Amber (soft tissues encased in hardened tree resin).

The identification of a fossil site as a Konservat-Lagerstätte may be based on a number of different factors which constitute "exceptional preservation". These may include the completeness of specimens, soft tissue preservation, fine-scale detail, taxonomic richness, distinctive taphonomic pathways (often multiple at the same site), the extent of the fossil layer in time and space, and particular sediment facies encouraging preservation.^[5]

Notable Lagerstätten

The world's major Lagerstätten include:

Precambrian

Site(s)	Age	Location	Significance	Notable fossils/organisms
Nonesuch Formation	1083-1070 Ma	Michigan, USA	An oxygenated Mesoproterozoic lake^[6] containing exceptionally preserved limnic microbes.^[7]
Lakhanda Lagerstätte	1030-1000 Ma	Uchur-Maya Depression, Russia	A site preserving a Mesoproterozoic community dominated by anaerobic bacteria.^[8] The lagerstätte contains evidence of trophic interactions from the Boring Billion.^[9]^[10]
Bitter Springs	1000–850 Ma	South Australia	Preserved fossils include cyanobacteria microfossils.	An early Stromatolite
Diabaig Formation	994 ± 48 Ma^[11]	Scotland	A freshwater environment preserving phosphatic microfossils,^[12] which represent some of the oldest known non-marine eukaryotes.^[13]
Dolores Creek Formation	950 Ma	Yukon, Canada	An Early Tonian site containing pyritised macroalgal fossils.^[14]
Chichkan Lagerstätte	775 Ma	Kazakhstan	A site from the transition between the prokaryote-dominated biota of the Early Neoproterozoic and the eukaryote-dominated biota of the Late Neoproterozoic and Phanerozoic.^[15]
Doushantuo Formation	600–555 Ma	Guizhou Province, China	Spans the poorly understood interval between the end of the Cryogenian period and the late Ediacaran Avalon explosion.	An Ediacaran embryo-like fossil
Portfjeld Formation	570 Ma	North Greenland	A Middle Ediacaran biota from the continent of Laurentia exhibiting Doushantuo-type preservation.^[16]
Mistaken Point	565 Ma	Newfoundland, Canada	This site contains one of the most diverse and well-preserved collections of Precambrian fossils.	Frondose ediacaran organisms
Itajai Biota	563 Ma	Brazil	An Ediacaran lagerstatte preserved by volcanism.^[17]
Ediacara Hills	555 Ma	South Australia	The type location the Ediacaran period, and has preserved a significant amount of fossils from that time.	Spriggina floundersi, a worm-like organism
Shibantan Lagerstätte	551-543 Ma	Hubei, China	A terminal Ediacaran fossil assemblage preserving life forms living just before the Proterozoic-Phanerozoic transition.^[18]
Gaojiashan Lagerstätte	551-541 Ma	Shaanxi, China	A lagerstätte documenting tube growth patterns of Cloudina.^[19]
Jiucheng Member	551-543 Ma	Yunnan, China	A latest Ediacaran macrofossil biota dominated by giant, unbranching thallophytes.^[20]
Khatyspyt Lagerstätte	544 Ma	Yakutia, Russia	A Late Ediacaran lagerstätte preserving an Avalon-type biota.^[21]
Bernashivka open pit	? (Upper Vendian)	Vinnytsia Oblast, Ukraine	A Late Ediacaran lagerstätte with numerous soft-bodied animals, algae, microfossils, bacteria, and fungi, comprising a number of different geological formations.^[22]

Cambrian

Site(s)	Age	Location	Significance	Notable fossils/organisms
Zhangjiagou	Fortunian	Shaanxi, China	A lagerstätte from the earliest Cambrian notable for its fossils of cnidarians,^[23] cycloneuralians,^[24]^[25] and the basal ecdysozoan Saccorhytus coronarius.^[26]	Life restoration of Saccorhytus as it may have been seen through the lens of a microscope, living among grains of sand.
Xiaoshiba Lagerstätte	Cambrian Stage 2	Yunnan, China	A site known for its detailed preservation of Early Cambrian macroalgae.^[27]
Maotianshan Shales (Chengjiang)	518 Ma	Yunnan, China	The preservation of an extremely diverse faunal assemblage renders the Maotianshan shale the world's most important for understanding the evolution of early multi-cellular life.	Haikouichthys, a primitive craniate
Qingjiang biota	518 Ma	Hubei, China	This site is particularly notable due to both the large proportion of new taxa represented (approximately 53% of the specimens), and the notable volume of soft-body tissue preservation.	Duplapex, a hymenocarine arthropod
Sirius Passet	523-518 Ma	Greenland	A site known for its fauna, and that they were most likely preserved by a death mask. It is a part of the larger Buen Formation, and has a fauna similar to the Maotianshan shales.	Kerygmachela, a dinocaridid-like lobopodian
Poleta Formation	519-518 Ma	Nevada, USA	The middle member of the formation preserves the Indian Springs Lagerstätte, one of the oldest such sites from former Laurentia. This site preserves a diversity of mineralized organisms such as trilobites and brachiopods, but also non-mineralized remains such as sponges, algae, and soft-bodied arthropods.^[28]	The trilobite Nevadia
Sinsk Algal Lens	518 Ma	Yakutia, Russia	One of the oldest known Cambrian lagerstätten. The fauna of this site is unique, as it seems that they were adapted to living in dysaerobic conditions.^[29]	A reconstruction of the large Lobopodian Siberion
Cranbrook Lagerstätte	Cambrian Stage 4	British Columbia	One of the oldest Burgess Shale-type biotas of North America.^[30]
Tatelt Formation	515 Ma	High Atlas, Morocco	A layer in this formation has produced some of the most well-preserved trilobites ever discovered, with preserved internal organs, feeding structures, and articulated appendages. The trilobites were likely rapidly buried and preserved by a volcanic eruption.^[31]	The trilobite Gigoutella mauretanica
Emu Bay Shale	513 Ma	South Australia	Noted soft tissue mineralization, most often of blocky apatite or fibrous calcium carbonate, including the oldest phosphatized muscle tissue.	Oestokerkus, a megacheiran arthropod
Parker Slate	513-511 Ma	Vermont, US	A Burgess Shale-type biota with rare but exceptionally preserved soft-bodied animals. The earliest Burgess Shale-type biota to be described, being documneted 25 years before the Burgess Shale itself.^[32]
Kaili Formation	513–501 Ma	Guizhou, China	The middle part of the Kaili Formation, the Oryctocephalus indicus Zone, contains a Burgess Shale-type lagerstätte with many well-preserved fossils known collectively as the Kaili Biota.	Pararotadiscus a cambroernid animal
Murero Lagerstätte	511-503 Ma	Spain	Thanks to the paleontological content, mainly trilobites, fourteen biozones have been established, the most precise biozonation for this time interval in the world. It also records in detail the so-called Valdemiedes event, the mass extinction episode at the end of the Lower Cambrian.^[33]	Eccaparadoxides trilobites showing sexual dimorphism
Blackberry Hill	~510–500 Ma	Central Wisconsin, US	This site preserves some of the oldest evidence of multicellular life walking out of the ocean, and onto dry land (in the form of large mollusks and euthycarcinoid arthropods). Other notable fossils include stranded scyphozoans, and some of the oldest true crustaceans (in the form of phyllocarids).	Mosineia, a euthycarcinoid arthropod
Henson Gletscher Formation	Wuliuan	North Greenland	A phosphatised lagerstätte preserving hatching priapulid larvae, pentastomids^[34] and abundant bradoriid and phosphatocopid arthropods.^[35]
Burgess Shale	508 Ma	British Columbia, Canada	One of the most famous fossil localities in the world. It is famous for the exceptional preservation of the soft parts of its fossils. At 508 million years old (middle Cambrian), it is one of the earliest fossil beds containing soft-part imprints.	Anomalocaris, a predatory radiodont
Spence Shale	507 Ma	Northeastern Utah, Southeastern Idaho, US	A site known for its abundant Cambrian trilobites and the preservation of Burgess Shale-type fossils. The type locality for this site is Spence Gulch in southeastern Idaho.	Hyoliths, most likely lophophorates
Linyi Lagerstätte	504 Ma	Shandong, China	A lagerstätte recognised for its exceptional preservation of arthropod limbs, intestines, and eyes.^[36]	Restoration of the Linyi Lagerstätte
Ravens Throat River Lagerstätte	Drumian	Northwest Territories	A Burgess Shale-type biota coeval in age with the more famous Wheeler Shale and Marjum Formation.^[37]
Wheeler Shale (House Range)	504 Ma	Western Utah, US	A world-famous locality known for its prolific agnostid and Elrathia kingii trilobite remains. Varied soft bodied organisms are also locally preserved, including Naraoia, Wiwaxia and Hallucigenia.	Elrathia sp. trilobites
Marjum Formation	502 Ma	Western Utah, US	A site known for its occasional preservation of soft-bodied tissue, and diverse assemblage.	Branchiocaris a pelagic hymenocarine
Weeks Formation	500 Ma	Western Utah, US	A site that is dominated by trilobites and brachiopods, but also comprising various soft-bodied organisms, such as Falcatamacaris.	Beckwithia, an aglaspidid trilobite relative
Kinnekulle Orsten and Alum Shale	500 Ma	Sweden	The Orsten sites reveals the oldest well-documented benthic meiofauna in the fossil record. Fossils such as microfossils of arthropods like free-living pentastomids are known. Multiple "Orsten-type" lagerstätten are also known from other countries.	Cambropachycope, a stem-group mandibulate arthropod

Ordovician

Site(s)	Age	Location	Significance	Notable fossils/organisms
Fezouata Formation	about 485 Ma	Draa Valley, Morocco	It was deposited in a marine environment, and is known for its exceptionally preserved fossils, filling an important preservational window beyond the earlier and more common Cambrian Burgess shale-type deposits.	Aegirocassis, a giant hurdiid radiodont
Cabrieres biota	Floian	Montagne Noire, France	A polar marine ecosystem from the Early Ordovician that likely served as a refuge from the high temperatures of the epoch.^[38]
Liexi fauna	About 470 Ma (early-middle Floian)	Hunan Province, China	Preserves Early Ordovician fauna with soft tissue, includes not only Cambrian relics but also taxa originated during Ordovician.^[39]
Castle Bank	About 461 Ma	Llandrindod Wells, Wales	A unique environment deposited during the middle Ordovician that possibly shows iconic groups from Cambrian lagerstättes, like Opabiniids and Megacheirans, survived for longer than what was thought.	Diagram of Mieridduryn, an opabiniid-like panarthropod
Douglas Dam Member^[40]	460 Ma	Tennessee, US	Low-diversity assemblage of arthropod fossils, which are preserved well because of volcanic ash.	Chasmataspis, a horseshoe crab-like chasmataspid arthropod
Winneshiek Shale	460 Ma	Decorah, Iowa	A Middle Ordovician site confined to a large impact Crater that is known for exceptionally exquisite preservation of conodonts, bivalved arthropods, and the earliest eurypterids in the fossil record.^[41]	Pentecopterus, the oldest known eurypterid
Beecher's Trilobite Bed	460? Ma	New York, US	Noted exceptionally preserved trilobites with soft tissue preserved by pyrite replacement. Pyritisation allows the use of X-rays to study fine detail of preserved soft body parts.	A pair of Triarthrus trilobites with pyritized soft-tissue
Harding Sandstone^[40]	? (Sandbian)	Colorado, US	Although preservation is not excellent, this lagoonal site provides early vertebrate fossils such as Astraspis and Eriptychius.	Eriptychius, an early agnathan
Walcott-Rust Quarry	about 455? Ma	New York, US	This site is an excellent example of an obrution (rapid burial or "smothered") Lagerstätte. Unique preservation of trilobite appendages resulted from early cementation of the surrounding rock and spar filling of the interior cavity of the appendages.	Isotelus gigas, a large asaphid trilobite
Big hill Lagerstätte	about 450? Mya	Michigan, US	A site known for its preservation of soft-bodied medusae (jellyfish), as well as linguloid brachiopods, algae, and arthropods (namely chasmataspidids, leperditid ostracods, and eurypterids).	Hoplitaspis, chasmataspid arthropod
Brechin Lagerstätte	450 Ma	Ontario, Canada	Known for preserving one of the most diverse crinoid fauna of the Katian.^[42]	Tomlinsonus, a marrellomorph
Soom Shale	450? Ma	South Africa	Known for its remarkable preservation of soft-tissue in fossil material. Deposited in still waters, the unit lacks bioturbation, perhaps indicating anoxic conditions.	Promissum, a conodont known from rare soft-tissues
Tafilalt Biota	? (Sandbian-middle Katian)	Tafilalt, Morocco	Known from range of non-biomineralised and soft-bodied organisms in polar environment.^[43]	"Eldonia" barvera, an eldoniid
Cat Head Member^[40]	? (middle Katian)	Manitoba, Canada	Fossils like algae, conulariids and trilobites are known from this site.
Georgian Bay Formation^[40]	449-445.6 Ma	Manitoulin, Canada	Low-diversity assemblage of arthropod fossils.	Orcanopterus, a middle-sized waeringopterid eurypterid
William Lake (Stony Mountain Formation)^[40]	445 Ma	Manitoba, Canada	Well-preserved fossils like jellyfish, xiphosurans, sea spiders are known from this site, it is important since many of the fossils are unknown in other Ordovician sites.	Lunataspis aurora, an early xiphosuran known in two sites
Airport Cove^[40]	445 Ma	Manitoba, Canada	Fossils like eurypterids, algae and xiphosurans are preserved in this site.	Lunataspis aurora, an early xiphosuran known in two sites

Silurian

Site(s)	Age	Location	Significance	Notable fossils/organisms
Kalana Lagerstätte	~440 Ma Aeronian	Estonia	Known for well preserved fossils of algae and crinoids,^[44] along with an osteostracan fossilised via an extremely unusual carbonaceous mode of preservation that was previously unknown among vertebrates.^[45]	Kalanaspis, the earliest known osteostracan
Chongqing Lagerstätte (Huixingshao Formation)^[46]	436 Ma	Chongqing, China	This site preserved complete fossils of earliest jawed vertebrates, as well as some galeaspids and eurypterids.	Xiushanosteus, the earliest placoderm
Waukesha Biota (Brandon Bridge Formation)	~435 Ma Early Silurian	Wisconsin, US	Well-studied site known for the exceptional preservation of its diverse, soft-bodied and lightly skeletonized fauna, includes many major taxa found nowhere else in strata of similar age. It was one of the first fossil sites with soft bodied preservation known to science.	Parioscorpio, an enigmatic arthropod
Herefordshire Lagerstätte (Coalbrookdale Formation)	~430 Ma	Herefordshire, UK	Known for the well-preserved fossils of various invertebrate animals many of which are in their three-dimensional structures. Fossils are preserved within volcanic ash, because of that sometimes this site has been compared to Pompeii.^[47] Some of the fossils are regarded as earliest evidences and evolutionary origin of some of the major groups of modern animals.	Offacolus, a euchelicerate
Eramosa Lagerstätte	~425 Ma	Ontario & New York (state)	Known for preservation of both hard and soft bodied organisms in great detail, including early scorpions, eurypterids, agnathan vertebrates, and several other species.	Eramoscorpius, an early scorpion
Bertie Group	422.9-416 Ma	Ontario & New York State	This limestone have produced thousands of fossil eurypterids, such as giant Acutiramus and well-known Eurypterus, as well as other fauna like scorpions and fish.	Nerepisacanthus, an acanthodian
Tonoloway Formation^[48]	~420 Ma	Pennsylvania, US	Known from exceptionally preserved mass assemblage of Eurypterus, the most abundant eurypterid in the fossil record.	Reconstruction of Eurypterus
Rochester Shale	415 Ma^[49]	New York, US and Ontario, Canada	Echinoderms (such as crinoids) and trilobites are known from Lewiston Member in this shale.	Arctinurus, one of the trilobites from this site

Devonian

Site(s)	Age	Location	Significance	Notable fossils/organisms
Rhynie chert	400 Ma	Scotland, UK	The Rhynie chert contains exceptionally preserved plant, fungus, lichen and animal material (euthycarcinoids, branchiopods, arachnids, hexapods, etc) preserved in place by an overlying volcanic deposit and hot springs. As well as one of the first known fully terrestrial ecosystems.	Asteroxylon, an early vascular plant related to lycopods
Waxweiler Lagerstätte (Klerf Formation)	409-392 Ma	Eifel, Germany	Waxweiler Lagerstätte is known from well-preserved fossils of chelicerates, giant claw of Jaekelopterus rhenaniae shows the largest arthropod ever known.	Jaekelopterus, around 2.5 m-long eurypterid
Heckelmann Mill	395 Ma	Rhineland-Palatinate, Germany	Heckelmann Mill preserves well preserved rhinocaridid archaeostracan phyllocarids,^[50] along with exceptionally abundant crinoid holdfasts from the late Emsian.^[51]
Hunsrück Slates (Bundenbach)	390 Ma	Rheinland-Pfalz, Germany	The Hunsrück slates are one of the few marine Devonian lagerstätte having soft tissue preservation, and in many cases fossils are coated by a pyritic surface layer.	Schinderhannes bartelsi, the youngest known radiodont
Gogo Formation	380 Ma (Frasnian)	Western Australia	The fossils of the Gogo Formation display three-dimensional soft-tissue preservation of tissues as fragile as nerves and embryos with umbilical cords. Over fifty species of fish have been described from the formation, and arthropods.	Materpiscis, a ptyctodontid placoderm fish that is the oldest vertebrate known to give live birth
Miguasha National Park (Escuminac Formation)	370 Ma	Québec, Canada	Some of the fish, fauna, and spore fossils found at Miguasha are rare and ancient species. For example, Eusthenopteron is sarcopterygian that shares characters with early tetrapods.	Fossil of lungfish Scaumenacia and antiarch placoderm Bothriolepis
Kowala Lagerstätte	~368 Ma	Świętokrzyskie Voivodeship, Poland	A Late Devonian site known for its fossils of non-biomineralised algae and arthropods.^[52]
Maïder Basin	368 Ma (for Thylacocephalan Layer)	Anti-Atlas, Morocco	Thylacocephalan Layer and Hangenberg Black Shale in this basin provides well-preserved fossils of Famennian fauna, including chondrichthyans and placoderms that preserved soft tissues.^[53]	Amazichthys, pelagic placoderm with preserved soft tissue
Strud^[54]	? (Late Famennian)	Namur Province, Belgium	Mainly juvenile placoderms are known, suggesting this site would be nursery site of placoderms.^[55] Various biota like tetrapods, arthropods and plants are also known, Strudiella from this site may be the earliest insect, but its affinity is disputed.	Strudops, the earliest known notostracan.
Canowindra, New South Wales (Mandagery Sandstone)	360 Ma	Australia	An accidentally discovered lagerstätte known for its exceptional preservation of Sarcopterygian and Placoderm fish.	Mandageria, a lobe-finned fish that is the NSW State Fossil Emblem in Australia
Waterloo Farm Lagerstätte (Witpoort Formation)	360 Ma	South Africa	Important site that providing the only record of a high latitude (near polar) coastal ecosystem, overturning numerous assumptions about high latitude conditions during the latest Devonian.	Priscomyzon, the oldest known genus of lamprey

Carboniferous

Site(s)	Age	Location	Significance	Notable fossils/organisms
Granton Shrimp Bed	? (Dinantian)	Firth of Forth, Scotland	Dominated by well-preserved crustacean fossils, this site provided first body fossil of Clydagnathus which solved long-lasted mystery of conodont fossils.
East Kirkton Quarry^[56]	335 Ma	West Lothian, Scotland	This site has produced numerous well-preserved fossils of early tetrapods like temnospondyls or reptiliomorphs, and large arthropods like scorpions or eurypterids.	Silvanerpeton, a possible reptiliomorph
Bear Gulch Limestone	324 Ma	Montana, US	A limestone-rich geological lens in central Montana. It is renowned for its unusual and ecologically diverse fossil composition of chondrichthyans, the group of cartilaginous fish containing modern sharks, rays, and chimaeras. Other animals like brachiopods, ray finned fish, arthropods, and the possible mollusk Typhloesus are also known from the site.	Falcatus, a holocephalian which males had large fin spine
Wamsutta Formation	~320-318 Ma	Massachusetts	A subhumid alluvian fan deposit that preserves ichnofossils, plants, invertebrates, and vertebrates.^[57]
Bickershaw^[58]	? (Langsettian)	Lancashire, England	This locality contains exceptionally preserved fossils within nodules. Arthropods have greater diversity, many of which are aquatic ones that lived in brackish environment.	Valloisella, a xiphosuran
Joggins Fossil Cliffs (Joggins Formation)	315 Ma	Nova Scotia, Canada	A fossil site that preserves a diverse terrestrial ecosystem consisting of plants like lycopsids, giant arthropods, fish, and the oldest known sauropsid, Hylonomus.	Hylonomus, the oldest known sauropsid in the fossil record
Castlecomer fauna	315-307	Kilkenny, Ireland	A konservat-lagerstätte with a high number of well-preserved spinicaudatan clam shrimp.^[59]^[60]
Linton Diamond Coal Mine^[61]^[62]	310 Ma	Ohio, US	A site known for its number of prehistoric tetrapods, like the lepospondyl Diceratosaurus.^[63]	The lepospondyl tetrapod Diceratosaurus
Mazon Creek	310 Ma	Illinois, US	A conservation lagerstätte found near Morris, in Grundy County, Illinois. The fossils from this site are preserved in ironstone concretions with exceptional detail. The fossils were preserved in a large delta system that covered much of the area. The state fossil of Illinois, the enigmatic animal Tullimonstrum, is only known from these deposits.	Tullimonstrum, an enigmatic animal
Buckhorn Asphalt Quarry	~310 Ma	Oklahoma, USA	A quarry of the Boggy Formation known for its exceptionally rich orthocerid assemblage.^[64]
Kinney Brick Quarry (Atrasado Formation)	around 307 Ma	New Mexico, US	This site is known from rich fish fossils with preserved soft tissues, that lived in lagoonal environment. Dozens of fish genera are known, ranging from chondrichthyans like ctenacanths and hybodonts, to actinopterygians and sarcopterygians.^[65]	Dracopristis, a ctenacanth
Campáleo Outcrop	Gzhelian	Santa Catarina, Brazil	A fungal and palynological lagerstätte from Gondwana during Late Palaeozoic Ice Age.^[66]
Montceau-les-Mines	300 Ma	France	Exceptional preservation of Late Carboniferous fossil biota are known, including various vertebrates and arthropods, as well as plants.^[67]^[68]	Idmonarachne, an arachnid that is related to spiders
Hamilton Quarry	300 Ma	Kansas, US	This site is known for its diverse assemblage of unusually well-preserved marine, euryhaline, freshwater, flying, and terrestrial fossils (invertebrates, vertebrates, and plants). This extraordinary mix of fossils suggests it was once an estuary.	Spinoaequalis, a semi-aquatic diapsid
Carrizo Arroyo	? (Latest Gzhelian to earliest Asselian)	New Mexico, US	This site is known from exceptional preservation of arthropod fossils, mainly insects.^[69]

Permian

Site(s)	Age	Location	Significance	Notable fossils/organisms
Taiyuan Formation	298 Ma	Inner Mongolia, China	Known from exceptionally well-preserved plant fossils in volcanic ash.^[70]^[71]
Meisenheim Formation^[72]	?(Asselian to early Sakmarian)^[73]	Lebach, Germany	This site is well-known for the rich occurrence of fauna lived in large freshwater lakes, including fish, temnospondyls, insects and others.	Lebachacanthus, a large-sized xenacanth
Franchesse	292 Ma	Massif Central, France	A Sakmarian seymouriamorph lagerstätte from the Bourbon l'Archambault Basin in the French Massif Central containing hundreds of complete seymouriamorph specimens.^[74]
Chemnitz petrified forest	291 Ma	Saxony, Germany	A petrified forest in Germany that is composed of Arthropitys bistriata, a type of Calamites, giant horsetails that are ancestors of modern horsetails, found on this location with never seen multiple branches. Many more plants and animals from this excavation are still in an ongoing research.^[75]	Large trunks of Arthropitys at Chemnitz
Mangrullo Formation	about 285–275 Ma (Artinskian)	Uruguay	This site is known for its abundant mesosaur fossils. It also contains the oldest known konservat-lagerstätte in South America, as well as the oldest known fossils of amniote embryos.^[76]	Stereosternum, an aquatic mesosaurid reptile
Chekarda (Koshelevka Formation)	about 283–273 Ma	Perm, Russia	Over 260 species of insect species are described from this site as well as diverse taxa of plants, making it one of the most important Permian konservat-lagerstätten.^[77]	Tshekardocoleidae beetle larva
Toploje Member	273-264 Ma	Prince Charles Mountains, Antarctica	This site preserves a high-latitude fauna in exceptional position before the large extinctions that happened later in the Permian.^[78]
Onder Karoo	266.9–264.28 Ma	Karoo Basin, South Africa	A high latitude, cool-temperate lacustrine ecosystem preserving detailed plant and insect fossils.^[79]
Sakamena Group^[80]	260–247 Ma	Madagascar	Lower Sakamena Formation (Permian) and Middle Sakamena Formation (Triassic) contain fossils of animals lived around wetland environment, such as semi-aquatic and gliding neodiapsids.	Claudiosaurus, aquatic neodiapsid reptile
Kupferschiefer	259–255 Ma	Central Europe	This site deposited in an open marine and shallow marine environment provides fossils of reptiles as well as many fish.	Weigeltisaurus, gliding weigeltisaurid reptile
Huopu Lagerstätte	~255 Ma	Guizhou, China	A plant fossil site documenting floral dynamics between the end-Guadalupian and end-Permian extinction events.^[81]

Triassic

Site(s)	Age	Location	Significance	Notable fossils/organisms
Guiyang biota^[82]	250.8 Ma	Guizhou Province, China	The oldest known Mesozoic lagerstätte (Dienerian). It preserves taxa belonging to 12 classes and 19 orders, including several species of fish.	Teffichthys elegans, a perleidiform ray-finned fish .
Paris biota^[83]	~249 Ma	Idaho, Nevada, USA	This earliest Spathian aged assemblage preserves fossils belonging to 7 phyla and 20 orders, combining Paleozoic groups (e.g. leptomitid protomonaxonid sponges otherwise known from the early Paleozoic) with members of the Modern evolutionary fauna (e.g. gladius-bearing coleoids).	Ankitokazocaris triassica is a thylacocephalan from the Paris biota
Jialingjiang Formation^[84]	249.2–247.2 Ma	Hubei Province, China	This site preserved aquatic reptiles soon after Permian extinction. Hupehsuchians are exclusively known from here, and already got unique ecology like filter feeding.	Eretmorhipis, a hupehsuchian that potentially had platypus-like ecology
Nanlinghu Formation^[84]	248 Ma	Anhui Province, China	This site provides important fossils to show early evolution of ichthyosauriforms.	Cartorhynchus, a primitive ichthyosauriform
Petropavlovka Formation	248 Ma	Orenburg Oblast, Russia	A site known for preserving oligochaetes, whose fossil record is extremely sparse.^[85]
Zarzaïtine Formation	Olenekian-Anisian	In Amenas, Algeria	A site with a high number of exceptionally well-preserved temnospondyl specimens, indicating of a seasonal climate with sudden droughts, with a freshwater ecosystem that could rapidly turn into a sebkha.^[86]
Luoping Biota (Guanling Formation)^[87]	~247-245 Ma	Yunnan, China	Various marine animals are preserved in this site, showing how marine ecosystem recovered after Permian extinction.^[88]	Atopodentatus, a herbivorous marine reptile that possessed a hammer-shaped head.
Grès à Voltzia	245 Ma	France	A fossil site remarkable for its detailed myriapod specimens.^[89] It also contains the earliest known aphid fossils.^[90]
Fossil Hill Member	? (Anisian)	Nevada, US	One of many Anisian marine lagerstatte, the Fossil Hill Member represents an open-ocean environment with a well-preserved fauna largely dominated almost entirely by ichthyosaurs.^[88]	The large ichthyosaur Thalattoarchon
Vossenveld Formation	? (Anisian)	Winterswijk,Netherlands	An exposure of this Muschelkalk formation in the Winterswijk quarry has a diverse assemblage of well-preserved marine reptiles, amphibians, fishes, and plants. It is the only marginal marine assemblage recorded from the earlier Triassic.^[88]
Strelovec Formation	? (Anisian)	Slovenia	A formation with well-preserved Triassic horseshoe crabs.^[91]
Saharonim Formation	Late Anisian/Lower Ladinian	Southern District, Israel	One brachiopod-dominated horizon of this formation documents the rapid burial of a community of exclusively juvenile Coenothyris brachiopods and ten different bivalve genera.^[92]
Besano Formation^[84]	242 Ma	Alps, Italy and Switzerland	This formation is designated as a World Heritage Site, as it is famous for its preservation of Middle Triassic marine life including fish and aquatic reptiles.^[88]^[93]	Tanystropheus, a long-necked archosauromorph
Xingyi biota (Zhuganpo Formation)^[84]	? (Upper Ladinian - Lower Carnian)	Guizhou and Yunnan, China	Previously considered as part of Falang Formation, this site yields many articulated skeletons of marine reptiles, as well as fish and invertebrates.	Keichousaurus, common sauropterygian from this site
Guanling biota (Xiaowa Formation)^[84]	? (Carnian)	Guizhou, China	Like Xingi Biota, this site also yields well-preserved marine fauna, especially many species of thalattosaurs are known.	Odontochelys, an early pantestudine (relative of turtles)
Polzberg	233 Ma	Austria	A site from the Reingrabener Schiefer known for exceptional preservation of bromalites^[94] and of cartilage,^[95] deposited during the Carnian Pluvial Event.^[96]	Fossil invertebrates from Polzberg
Madygen Formation	230 Ma	Kyrgyzstan	The Madygen Formation is renowned for the preservation of more than 20,000 fossil insects, making it one of the richest Triassic lagerstätten in the world. Other vertebrate fossils as fish, amphibians, reptiles and synapsids have been recovered from the formation too, as well as minor fossil flora.	Longisquama, a bizarre reptile with long growths protruding from its back
Cow Branch Formation	230 Ma	Virginia, US	This site preserves a wide variety of organisms (including Fish, reptiles, arachnids, and insects).	Mecistotrachelos, a gliding reptile distantly related to archosauromorphs, like crocodylians and dinosaurs
Alakir Çay	? (Norian)	southwest Turkey	A konservat-lagerstätte with exceptionally well-preserved Triassic corals, retaining much of their original aragonite skeletons.^[97]

Jurassic

Site(s)	Age	Location	Significance	Notable fossils/organisms
Osteno (Moltrasio Formation)^[98]	196-188 Ma	Italy	Several kinds of marine biota such as fish, crustaceans, cephalopods polychaetes, and nematodes have been recovered. This site is the only fossil deposit in Italy in which soft bodies are preserved other than Monte Bolca.	Ecological reconstruction in the site, showing thylacocephalan Ostenocaris and cephalopod Ostenoteuthis
Ya Ha Tinda	183 Ma	Alberta, Canada	A fossil site notable for containing abundant and extremely well-preserved vampire squid, being the largest concentration of vampire squid fossils outside the Tethys Ocean,^[99] and for being deposited during the Toarcian Oceanic Anoxic Event (TOAE).^[100]^[101]^[102]
Strawberry Bank	183 Ma	Somerset, England	A site from the TOAE documenting marine life during the recovery from the Triassic-Jurassic extinction event as well as the turmoil of the TOAE.^[103] The oldest pseudoplanktonic barnacles in the fossil record,^[104] near-complete ichthyosaur skeletons,^[105] and evidence of ichthyosaur niche partitioning are preserved at this site.^[106]
Holzmaden/Posidonia Shale	183 Ma	Württemberg, Germany	The Sachrang member is among the most important formations of the Toarcian boundary, due to the concentrations of exceptionally well-preserved complete skeletons of fossil marine fish and reptiles. It was also deposited during the TOAE.^[107]^[108]	A specimen of the ichthyosaur Stenopterygius crassicostatus with preserved young
Cabeço da Ladeira	Late Bajocian	Portugal	A site known for exquisite preservation of microbial mats in a tidal flat.^[109]
Monte Fallano	? (Bajocian-Bathonian)	Campania, Italy	This Plattenkalk preserves fossils of terrestrial plants, crustaceans and fish.^[110]
Christian Malford	Callovian	Wiltshire, England	A site in the Oxford Clay Formation which preserves exceptionally detailed coleoid fossils.^[111]
Mesa Chelonia^[112]	164.6 Ma	Shanshan County, China	This site is notable because it contains a large turtle bonebed, containing specimens of the genus Annemys. This bonebed contains up to an estimated 36 turtles per square meter.
La Voulte-sur-Rhône	160 Mya	Ardèche, France	La Voulte-sur-Rhône, in the Ardèche region of southwestern France, offers paleontologists an outstanding view of an undisturbed paleoecosystem that was preserved in fine detail. Notable finds include retinal structures in the eyes of thylacocephalan arthropods, and fossilized relatives of the modern day vampire squid, like Vampyronassa rhodanica.	A rare well-preserved cephalopod, Rhomboteuthis
Shar Teeg Beds	160-145 Mya	Govi-Altay, Mongolia	Many of insect remains and some vertebrates like relatives of crocodilians are known from this site.^[113]
Karabastau Formation	155.7 Ma	Kazakhstan	This site is an important locality for insect fossils that has been studied since the early 20th century, alongside the rarer remains of vertebrates, including pterosaurs, salamanders, lizards and crocodiles.	Sordes, small pterosaur with visible soft-tissues preserved
Tiaojishan Formation	165-153 Ma	Liaoning Province, China	It is known for its exceptionally preserved fossils, including those of plants, insects and vertebrates. It is made up mainly of pyroclastic rock interspersed with basic volcanic and sedimentary rocks.	Anchiornis, small feathered anchiornithid dinosaur
La Casita Formation	Kimmeridgian	Coahuila, Mexico	A marine konzentrat-lagerstätte deposited in a hemipelagic mud bottom during dysoxic conditions.^[114]
Talbragar fossil site^[115]	151 Ma	New South Wales, Australia	This bed is part of Purlawaugh Formation, and provided fauna like fish and insects that lived around the lake.	Cavenderichthys, a teleost known from this site
Cleveland-Lloyd Dinosaur Quarry	150 Ma	Utah, US	Jurassic National Monument, at the site of the Cleveland-Lloyd Dinosaur Quarry, well known for containing the densest concentration of Jurassic dinosaur fossils ever found, is a paleontological site located near Cleveland, Utah, in the San Rafael Swell, a part of the geological layers known as the Morrison Formation. Up to 15,000 have been excavated from this site alone.	Allosaurus, the largest predator of the quarry
Canjuers Lagerstätte	150 Ma	France	This site shows a high amount of biodiversity, including reptiles, invertebrates, fish, and other organisms.	Pleurosaurus, an aquatic rhynchocephalian
Agardhfjellet Formation	150-140 Ma	Spitsbergen, Norway	The formation contains the Slottsmøya Member, a highly fossiliferous unit where many ichthyosaur and plesiosaur fossils have been found, as well as abundant and well preserved fossils of invertebrates.	Pliosaurus funkei, a large thalassophonean pliosaurid which was informally known as "Predator X"
Solnhofen Limestone	149-148 Ma	Bavaria, Germany	This site is unique as it preserves a rare assemblage of fossilized organisms, including highly detailed imprints of soft bodied organisms such as sea jellies. The most familiar fossils of the Solnhofen Plattenkalk include the early feathered theropod dinosaur Archaeopteryx preserved in such detail that they are among the most famous and most beautiful fossils in the world.	The Berlin Specimen of Archaeopteryx lithographica
Owadów–Brzezinki site	~148 Ma	Łódź Voivodeship, Poland	A marine deposit of the Kcynia Formation similar to the Solnhofen Formation, with large numbers of preserved insect remains, numerous marine invertebrates, and vertebrates including fishes, marine reptiles, and pterosaurs.^[116]^[117]	Fossil of the horseshoe crab Crenatolimulus darwini

Cretaceous

Site(s)	Age	Location	Significance	Notable fossils/organisms
Angeac-Charente bonebed	~141 Ma	Charente, France	A lagerstätte preserving both vertebrate and invertebrate fossils from the poorly represented Berriasian stage known for its taphonomic and sedimentological ‘frozen scenes’.^[118]
El Montsec (La Pedrera de Rúbies Formation)	~140-125 Ma	Catalonia, Spain	Known from exceptional preservation of biota such as plants, fish, insects, crustaceans and even some tetrapods.^[119]	Montsecosuchus, atoposaurid crocodylomorph
Lebanese amber	~130-125 Ma (Barremian)	Lebanon	Preserves a high diversity of insects from the Early Cretaceous, and is among the oldest known fossilized amber to contain a significant number of preserved organisms.^[120] Includes many of the oldest known members of modern insect groups, and many of the youngest known members for extinct insect groups.^[121]	Preserved larval lacewings (Tragichrysa ovoruptora)
Las Hoyas	about 125 Ma (Barremian)	Cuenca, Spain	The site is mostly known for its exquisitely preserved dinosaurs, especially enantiornithines. The lithology of the formation mostly consists of lacustarine limestone deposited in a freshwater wetland environment.	Concornis, an early enantiornithean
Yixian Formation	about 125–121 Ma (Barremian-Aptian)	Liaoning, China	The Yixian Formation is well known for its great diversity of well-preserved specimens and its feathered dinosaurs, such as the large tyrannosauroid Yutyrannus, the therizinosaur Beipiaosaurus, and various small birds, along with a selection of other dinosaurs, such as the iguanodontian Bolong, the sauropod Dongbeititan and the ceratopsian Psittacosaurus. Other biota included the troodontid Mei, the dromaeosaurid Tianyuraptor, and the compsognathid Sinosauropteryx.	Sinosauropteryx, the first non-avain dinosaur with evidence of feathers to have been recognized
Jiufotang Formation	about 122-119 Ma (Aptian)	Liaoning, China	This formation overlies the slightly older Yixian Formation and preserved very similar species, including a wide variety of dinosaurs such as the ceratopsian Psittacosaurus and the early bird Confuciusornis, both of which are also found in the Yixian Formation. Also notable are the very abundant specimens of the dromaeosaurid Microraptor, which is known from up to 300 specimens and is among the most common animals found here.	Microraptor had four wings and is believed to have evolved powered flight independently of true birds.
Khasurty Fossil Site	? (Aptian)	Buryatia, Russia	One of the largest fossil insect sites in northern Asia, with over 6000 fossilized insect specimens preserved in mudstones, representing over 16 orders and 130 families. Taxa have both Jurassic & Cretaceous affinities. Fossils of other invertebrates such as arachnids & crustaceans are also known, in addition to small plants and fragmentary vertebrate remains such as fish scales and bird feathers.^[122]
Shengjinkou Formation	about 120 Ma	Xinjiang, China	Part of the finds from this site consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. The site represented a nesting colony that storm floods had covered with mud. Dozens of individuals could be secured from a total that in 2014 was estimated to run into the many hundreds.	Hamititan, a large titanosaurian sauropod
Xiagou Formation	about 120–115? Ma	Gansu, China	This site is known outside the specialized world of Chinese geology as the site of a lagerstätte in which the fossils were preserved of Gansus yumenensis, the earliest true modern bird.	Gansus is among the earliest known modern birds
Paja Formation	130-113 Ma	Colombia	This site is famous for its vertebrate fossils and is the richest Mesozoic fossiliferous formation of Colombia. Several marine reptile fossils of plesiosaurs, pliosaurs, ichthyosauras and turtles have been described from the formation and it hosts the only dinosaur fossils described in the country to date; the titanosauriform sauropod Padillasaurus.	Desmatochelys, a protostegid and one of the oldest known turtle fossils
Koonwarra Fossil Bed^[123]	around 118-115 Ma	Victoria, Australia	This site is composed of mudstone sediment thought to have been laid down in a freshwater lake. Arthropods, fish and plant fossils are known from this site.	Tarwinia, a large sized stem-flea
Crato Formation	113 Ma	northeast Brazil	The Crato Formation earns the designation of lagerstätte due to an exceedingly well preserved and diverse fossil faunal assemblage. Some 25 species of fossil fishes are often found with stomach contents preserved, enabling paleontologists to study predator-prey relationships in this ecosystem. There are also fine examples of pterosaurs, reptiles and amphibians, invertebrates (particularly insects), and plants. Also known from this site is Ubirajara, the first non-avian dinosaur from the southern hemisphere with evidence of feathers. Additionally, the formation abounds with evidence of plant-insect interaction.^[124]	Tupandactylus, a fossil pterosaur that was preserved with feathers and other soft tissues intact
Amargosa Bed (Marizal Formation)	? (Aptian-Albian)	northeast Brazil	Fluvial site which preserved fish, crustacean and plant fossils.^[125]
Pietraroja Plattenkalk	113-110 Ma	Campania, Italy	A konservat-lagerstätte famous for its diverse and well-preserved fish and plant fossils. Also known from this formation is Scipionyx, one of Europe's most well-preserved dinosaurs.^[126]^[127]	Scipionyx, a small dinosaur with preserved internal organs.
Jinju Formation	112.4–106.5 Ma	South Korea	The Jinju Formation is notable for the post-Jehol Group insect assemblage,^[128] as well as other fauna such as isopods and fish.^[129]^[130] The site is also notable for its abundance and diversity of tetrapod trackways.^[131]	Coptoclavidae beetle larva preserved at this locality
Tlayúa Formation	110 Ma	Puebla, Mexico	A marine lagerstätte preserving Albian actinopterygians and lepidosaurs.^[132]	Cipactlichthys, example of fish from this site
Romualdo Formation	108–92 Ma	northeast Brazil	The Romualdo Formation is a part of the Santana Group and has provided a rich assemblage of fossils; flora, fish, arthropods insects, turtles, snakes, dinosaurs, such as Irritator, and pterosaurs, including the genus Thalassodromeus. The stratigraphic units of the group contained several feathers of birds, among those the first record of Mesozoic birds in Brazil.	Tapejara, a common pterosaur in Brazil during the Early Cretaceous
Muhi Quarry (El Doctor Formation)	? (Albian to Cenomanian, probably Late Albian)^[133]	Hidalgo, Mexico	While this site produced limestones for construction, rocks in that locality contain a diverse Cretaceous marine biota such as fish, ammonites and crustaceans.	Motlayoichthys, a member of Pachyrhizodontidae
Puy-Puy Lagerstätte	100.5 Ma	France	A paralic site preserving a variety of ichnofossils,^[134] along with some vertebrate remains.^[135] The site preserves evidence of plant-insect interaction.^[136]
Burmese amber	101-99 Ma (latest Albian/earliest Cenomanian)	Myanmar	More than 1,000 species of taxa have been described from ambers from Hukawng Valley. While it is important for understanding the evolution of biota, mainly insects, during the Cretaceous period, it is also extremely controversial by facing ethical issues due to its association with conflicts and labor conditions.	Oculudentavis, small-sized lizard
English Chalk	100-90 Ma (Cenomanian to Turonian)	England	Two subsections of England's famous chalk formation, the Grey Chalk Subgroup and the lower sections of the White Chalk Subgroup, yield three-dimensionally preserved fossils of marine fishes. This exquisite level of preservation is unlike fish fossils from other deposits from around the same time, which are only preserved as two-dimensional compression fossils.^[137]	Fossil of the fish Aulolepis typus
Haqel/Hjoula/al-Nammoura	95-94 Ma	Lebanon	Famous Lebanese konservat-lagerstätten of the Late Cretaceous (middle to late Cenomanian) age, which contain a well-preserved variety of different fossils. Small animals like shrimp, octopus, stingrays, and bony fishes are common finds at these sites. Some of the rarest fossils from this locality include those of octopuses.^[138]	Ichthyoceros, pycnodont actinopterygian
Komen Limestone	95-94 Ma	Komen, Slovenia	A Late Cenomanian locality in the Karst of Slovenia with a high diversity of articulated fossil fish, in addition to small reptiles and invertebrates.^[139]	The marine lizard Carsosaurus
Hesseltal Formation	94–93 Ma	Saxony & North Rhine-Westphalia, Germany	Deposited during the anoxic conditions of the Cenomanian-Turonian boundary event, this formation has a high number of well-preserved, articulated fish skeletons, in addition to exceptionally preserved ammonites with soft parts.^[140]^[141]	The billfish-like Rhamphoichthys
Vallecillo (Agua Nueva Formation)	94–92 Ma	Nuevo León, Mexico	The site is noted for its qualities as a konservat-lagerstätte, with notable finds including the plesiosaur Mauriciosaurus and the possible shark Aquilolamna.	Mauriciosaurus, a short-necked plesiosaur
Akrabou Formation (Gara Sbaa/Agoult & Goulmima)	? (Turonian)	Asfla, Morocco	Marine site known for exceptionally preserved, three-dimensional fish fossils.^[142]^[143]	Grandemarinus, an early gar
Orapa diamond mine	Turonian	Botswana	An insect lagerstätte known for being one of the few entomofaunas from southern Africa, containing a variety of insects,^[144] particularly beetles.^[145]^[146]
New Jersey amber	91-89 Ma	New Jersey, US	Turonian-aged amber from the Raritan & Magothy Formations of New Jersey, with a high diversity of well-preserved insects, plants and fungi.^[147]	Sphecomyrma, an early ant
Lower Idzików beds	87-86 Ma (Coniacian)	Lower Silesian Voivodeship, Poland	An exposure of these beds near Stary Waliszów contains a konzentrat-lagerstätte of numerous Cretaceous marine invertebrates in concretions, including decapods, molluscs and echinoderms, as well as well-preserved plant fossils that indicate a nearshore environment. Very well-preserved phosphatized decapod remains are known.^[148]
Smoky Hill Chalk	87–82 Ma	Kansas and Nebraska, US	A Cretaceous konservat-lagerstätte known primarily for its exceptionally well-preserved marine reptiles. Also known from this site are fossils of large bony fish such as Xiphactinus, mosasaurs, flying reptiles or pterosaurs (namely Pteranodon), flightless marine birds such as Hesperornis, and turtles.	Xiphactinus is famous for being found with another fish (Gillicus) preserved in its stomach.
Ingersoll Shale	85 Ma	Alabama, US	A Late Cretaceous (Santonian) informal geological unit in eastern Alabama. Fourteen theropod feathers assigned to birds and possibly dromaeosaurids have been recovered from the unit.
Sahel Alma	~84 Ma	Lebanon	A Late Cretaceous (Santonian) konservat-lagerstätte with similar excellent preservation of marine organisms as the nearby, older Sannine Lagerstätte, but in a deep-water environment. Includes a high number of well-preserved shark body fossils, in addition to cephalopods and deepwater arthropods.^[138]	Propristiophorus, an early sawshark
Auca Mahuevo	80 Ma	Patagonia, Argentina	A Cretaceous lagerstätte in the eroded badlands of the Patagonian province of Neuquén, Argentina. The sedimentary layers of the Anacleto Formation at Auca Mahuevo were deposited between 83.5 and 79.5 million years before the present and offers a view of a fossilized titanosaurid nesting site.	An egg from a titanosaurian sauropod
Ellisdale Fossil Site	79-76 Ma	New Jersey, US	A middle Campanian konzentrat-lagerstätte from the Marshalltown Formation with one of the most diverse Mesozoic vertebrate faunas of eastern North America, likely originating from a flood event. A high number of disarticulated bones of dinosaurs, fish, reptiles, amphibians, and small mammals is known, most of which are microfossils.^[149]
Coon Creek Formation	76.8-76.0 Ma^[150]	Tennessee and Mississippi, US	This late Campanian formation has some of the world's best-preserved remains of Cretaceous marine invertebrates (primarily mollusks and decapod crustaceans), with many retaining their original aragonitic shells and exoskeletons.^[150]^[151]	The fossil crab Avitelmessus grapsoides, which occurs in great numbers at Coon Creek
Baumberge Formation	~75-72 Ma	North Rhine-Westphalia, Germany	A Late Campanian formation in the Baumberge of Germany with a high number of articulated fossil fish remains, in addition to shark body fossils.^[140]^[152]	The paracanthopterygian fish Sphenocephalus
Nardò (Calcari di Melissano)^[153]	~72-70 Ma (upper Campanian-lower Maastrichtian)	Apulia, Italy	This site is especially famous for its limestones containing abundant fossil fish remains.	Nardovelifer, the earliest lampriform
Harrana (Muwaqqar Chalk Marl Formation)	66.5-66.1 Ma (Late Maastrichtian)^[154]	Jordan	Phosphatic deposits formed in this site are known to preserve vertebrate fossils with soft tissue, such as mosasaurs, plesiosaurs, sharks, bony fish, turtles and crocodylians.^[155]	Postredectes, an example of fish fossil from this site
Zhucheng (Wangshi Group)	66 Ma	Shandong, China	Zhucheng has been an important site for dinosaur excavation since 1960. The world's largest hadrosaurid fossil (Shantungosaurus) was found in Zhucheng in the 1980s. Other dinosaurs known from the area include the ceratopsian Zhuchengceratops (2010), the sauropod Zhuchengtitan (2017) and the theropod Zhuchengtyrannus (2011) which have all been described from deposits near and named after Zhucheng.	Zhuchengtyrannus, an Asian relative of the larger Tyrannosaurus
Tanis^[156]	66.0 Ma	North Dakota, US	Tanis is part of the heavily studied Hell Creek Formation, a group of rocks spanning four states in North America renowned for many significant fossil discoveries from the Upper Cretaceous and lower Paleocene. Tanis is a significant site because it appears to record the events from the first minutes until a few hours after the impact of the giant Chicxulub asteroid in extreme detail. This impact, which struck the Gulf of Mexico 66.043 million years ago, wiped out all non-avian dinosaurs and many other species (the so-called "K-Pg" or "K-T" extinction).	Fossil sturgeon from the Tanis site

Paleogene

Site(s)	Age	Location	Significance	Notable fossils/organisms
Baunekule Facies	64-63 Ma (Danian)	eastern Denmark	These facies of the Faxe Formation document an extremely well-preserved cold-water coral mound ecosystem dominated by Dendrophyllia corals, and also includes gastropods, tubeworms, bivalves, bryozoans and gastropods.^[157]
Tenejapa-Lacandón Formation	~63 Ma	Chiapas, Mexico	A formation with a high number of well-preserved fish fossils indicative of mass mortality events.^[158] One of the most important formations for documenting the recovery of ocean ecosystems in the wake of the K-Pg extinction, due to being deposited just a few million years after and being located only 500 kilometres (310 mi) away from the Chixculub impact site.^[159]	Eekaulostomus, Kelmejtubus, and Paleoserranus, three unique fish
Menat	60 Ma	Auvergne, France	A Palaeocene maar lake containing three-dimensional plant remains.^[160] It is particularly notable for preserving one of the oldest known bees.^[161]
Danata Formation	56-53 Ma	western Turkmenistan	Outcropping in the Kopet Dag range, this formation preserves numerous fossil fish from a northeastern arm of the Tethys Ocean during the Paleocene-Eocene thermal maximum. 38 taxa from 13 orders are known, the vast majority of which are acanthomorphs.^[162]^[163]	Eospinus, a bizarre tetraodontiform fish.
Fur Formation	55–53 Ma	Fur & Mors, Denmark	Preserves abundant fossil fish, insects, reptiles, birds and plants. The Fur Formation was deposited about 55 Ma, just after the Palaeocene-Eocene boundary, and its tropical or sub-tropical flora indicate that the climate after the Paleocene-Eocene Thermal Maximum was moderately warm (approximately 4-8 degrees warmer than today).	Tasbacka danica, immature sea turtle
London Clay	54–48 Ma	England, UK	Collected for close to 300 years, Plant fossils, especially seeds and fruits, are found in abundance. Some 350 named species of plant have been found, making the London Clay flora one of the world's most diverse for fossil seeds and fruits. The flora includes tropical taxa found in modern Asia, reflecting the much warmer climate of the early Eocene.	A Tynskya eocaena fossil
Eocene Okanagan Highlands	52 - 48 Ma	British Columbia, Canada & Washington, USA	Includes McAbee Fossil Beds, Princeton chert & Klondike Mountain Formation; Recognized as temperate/subtropical uplands right after the Paleocene–Eocene Thermal Maximum and spanning the Early Eocene Climatic Optimum, preserves highly detailed uplands lacustrine fauna and flora.	A Florissantia quilchenensis flower, an extinct malvaceous genus from western North America
Monte Solane	51-49 MA	Verona, Italy	Slightly older than the nearby, more well-known Monte Bolca site, the Monte Solane site also preserves numerous marine fish and plants, but documents an entirely different ecosystem that appears to be of a bathypelagic habitat, forming one of the few known lagerstätte to preserve a deep-sea ecosystem.^[164]
Green River Formation	50 Ma	Colorado/Utah/Wyoming, US	An Eocene aged site that is noted for the fish fauna preserved. Other fossils include the crocodilians, birds, and mammals.	Diplomystus and Knightia
Monte Bolca	50-49 Ma	Verona, Italy	A fossil site with specimens of fish and other organisms that are so highly preserved that their organs are often completely intact in fossil form, and even the skin color can sometimes be determined. It is assumed that mud at the site was low in oxygen, preventing both decay and the mixing action of scavengers from harming the fossils.	A complete Archaeophis proavus
Messel Formation	47 Ma	Hessen, Germany	This site has significant geological and scientific importance. Over 1000 species of plants and animals have been found at the site. After almost becoming a landfill, strong local resistance eventually stopped these plans and the Messel Pit was declared a UNESCO World Heritage Site on 9 December 1995. Significant scientific discoveries about the early evolution of mammals and birds are still being made at the Messel Pit, and the site has increasingly become a tourist site as well.	Masillamys sp., an ischyromyid rodent
Baltic amber	47-35 Ma (Lutetian to Priabonian)	Pomeranian Voivodeship, Poland & Kaliningrad Oblast, Russia	The largest amber deposit on Earth, this amber is part of the Prussian Formation, and preserves a high diversity of exceptionally well-preserved fossil invertebrates, plants, and small vertebrates that inhabited eastern Europe during the warmer, subtropical conditions of the middle Eocene. It is the largest world's single largest repository of fossil insects.^[165]^[166]^[167]^[168]	Yantarogekko, a gecko
Kishenehn Formation	46.2 Ma	Montana	A Middle Eocene site preserving exquisitely detailed insect specimens in oil shale.^[169]
Mahenge	46 Ma	Tanzania	A terrestrial Middle Eocene lagerstätte preserving plant and arthropod fossils.^[170]^[171]
Quercy Phosphorites Formation^[172]	45-25 Ma	Occitania, France	This site 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.	Xenomorphia resurrecta parasitic wasps preserved in fly pupae
Na Duong	Priabonian	Northern Vietnam	A Late Eocene site notable for its highly detailed coprolite preservation.^[173]
Bitterfeld amber	38-34 Ma	Saxony, Germany	One of three major Paleogene deposits of European amber, this deposit of the Cottbus Formation shares a similar biota to the Baltic Amber, indicating a concurrent formation, but appears to have a geologically distinct origin from Baltic amber.^[168]	Pseudoscorpions from the Bitterfeld amber
Rovno amber	38-34 Ma	Rivne Oblast, Ukraine & Gomel, Belarus	One of three major Paleogene deposits of European amber, this deposit of the Obukhov Formation preserves a high diversity of invertebrates, many of which are shared with the Baltic amber but others of which are unique. A drier habitat compared to the Baltic amber is suggested based on some of the insect taxa preserved.^[165]	Eocenomyrma ukrainica, an ant
Florissant Formation	34 Ma	Colorado, US	A late Eocene (Priabonian) aged site that is noted for the finely preserved plant and insect paleobiota. Fossils are preserved in diatom blooms of a lahar dammed lake system and the formation is noted for the petrified stumps of Sequoia affinis	A Palaeovespa florissantia
Canyon Ferry Reservoir	32.0 ± 0.1 Ma	Montana, US	A diverse Early Oligocene plant and insect fossil site.^[174]
Rauenberg	30 Ma	Baden-Württemberg, Germany	A marine fossil site with an Arctic-like invertebrate fauna and a Paratethyan vertebrate fauna displaying evidence of intermittent anoxia.^[175]
Sangtang Lagerstätte	~28 – 23 Ma	Guangxi, China	A section of the Late Oligocene Yongning Formation with one of the very few known Cenozoic assemblages of mummified plant fossils,^[176] including mummified fruits.^[177]^[178]^[179]
Enspel Lagerstätte	24.79-24.56 Ma	Rhineland-Palatinate, Germany	A Chattian maar deposit famous for its insect fossils.^[180]
Aix-en-Provence	~24 Ma	Provence, France	A terminal Oligocene brackish palaeoenvironment.^[181]

Neogene

Site(s)	Age	Location	Significance	Notable fossils/organisms
Dominican amber	30–10 Ma	Dominican Republic	Dominican amber differentiates itself from Baltic amber by being nearly always^{[citation needed]} transparent, and it has a higher number of fossil inclusions. This has enabled the detailed reconstruction of the ecosystem of a long-vanished tropical forest.^[182]	A larval Amblyomma tick preserved in amber.
Riversleigh	25–15 Ma	Queensland, Australia	This locality is recognised for the series of well preserved fossils deposited from the Late Oligocene to the Miocene. The fossiliferous limestone system is located near the Gregory River in the north-west of Queensland, an environment that was once a very wet rainforest that became more arid as the Gondwanan land masses separated and the Australian continent moved north.	Reconstruction of the diprotodont marsupial Nimbadon lavarackorum.
Foulden Maar	23 Ma	Otago, New Zealand	These layers of diatomite have preserved exceptional fossils of fish from the crater lake, and plants, spiders, and insects from the sub-tropical forest that developed around the crater,^[183] along with in situ pollen.^[184]	The teleost fish species Galaxias effusus.
Ebelsberg Formation	23-22 Ma	Upper Austria, Austria	This Aquitanian-aged konservat-lagerstätte records an exceptional fossil assemblage of an enormous number of plants, fish, marine mammals, and marine invertebrates from a section of the central Paratethys Sea.^[185]	Austromola, a sunfish
Chiapas amber	23-15 Ma	Chiapas, Mexico^[186]	As with other ambers, a wide variety of taxa have been found as inclusions including insects and other arthropods, as well as plant fragments and epiphyllous fungi.	The holotype specimen of the millipede Anbarrhacus adamantis.
Clarkia fossil beds	20-17 Ma	Idaho, US	The Clarkia fossil beds site is best known for its fossil leaves. Their preservation is exquisite; fresh leaves are unfossilized, and sometimes retain their fall colors before rapidly oxidizing in air. It has been reported that scientists have managed to isolate small amounts of ancient DNA from fossil leaves from this site. However, other scientists are skeptical of the validity of this reported occurrence of Miocene DNA.	A leaf fossil from the beds after being exposed to oxygen.
Barstow Formation	19–13.4 Ma	California, US	The sediments are fluvial and lacustrine in origin except for nine layers of rhyolitic tuff. It is well known for its abundant vertebrate fossils including bones, teeth and footprints. The formation is also renowned for the fossiliferous concretions in its upper member, which contain three-dimensionally preserved arthropods.	A fossilized footprint made by a camel.
Shanwang Formation	18-17 Ma	Shandong Province, China	Fossils have been found at this site in dozens of categories, representing over 600 separate species. Animal fossils include insects, fish, spiders, amphibians, reptiles, birds and mammals. Insect fossils have clear, intact veins. Some have retained beautiful colours.	Fossil of Lusorex
Morozaki Group^[187]	18-17 Ma	Aichi Prefecture, Japan	Known from well-preserved deep sea fauna including fish, starfish and arthropods like crabs, shrimps and giant amphiopods.
Sandelzhausen	16 Ma	Bavaria, Germany	A Middle Miocene vertebrate locality.^[188]
McGraths Flat	~16-11 Ma	NSW, Australia	Deposited in unusual conditions that record microscopic details of soft tissues and delicate structures. Fossil evidence of animals with soft bodies, unlike the bones of mammals and reptiles, is rare in Australia, and discoveries at McGraths' Flat have revealed unknown species of invertebrates such as insects and spiders.^[189]
Dolnja Stara	~15 Ma	Slovenia	A barnacle fossil site preserving barnacles shortly after settlement attached to mangrove leaves.^[190]
Pisco Formation	15-2 Ma	Arequipa & Ica, Peru	Several specialists consider the Pisco Formation one of the most important lagerstätten, based on the large amount of exceptionally preserved marine fossils, including sharks (most notably megalodon), penguins, whales, dolphins, birds, marine crocodiles and aquatic giant sloths.	Reconstruction of the macroraptorial stem-physeteroid whale Acrophyseter.
Hindon Maar	14.6 Ma	New Zealand	A maar preserving a Southern Hemisphere lake-forest ecosystem, including body fossils of plants, insects, fish, and birds,^[191] along with in situ pollen^[184] and coprolites of both fish and birds.^[191]
Ngorora Formation	13.3-9 Ma	Tanzania	The alkaline palaeolake deposits of the Ngorora Formation contains articulated fish fossils that died en masse from asphyxiation during episodic ash falls or from rapid acidification.^[192]	Megistotherium was a large sized hyaenodont mammal
Pi Gros	13 Ma	Catalonia, Spain	An ichnofossil lagerstätte containing annelid, mollusc, and sponge trace fossils. The fossil site no longer exists due to having been quarried for the construction of an industrial park.^[193]
Bullock Creek	12 Ma	Northern Territory, Australia	Among the fossils at the Bullock Creek site have been found complete marsupial crania with delicate structures intact. New significant taxa identified from the Bullock Creek mid Miocene include a new genus of crocodile, Baru (Baru darrowi), a primitive true kangaroo, Nambaroo, with high-crowned lophodont teeth; and a new species of giant horned tortoise, Meiolania. New marsupial lion, thylacine, and dasyurid material has also been recovered.
Tunjice	? (Middle Miocene)	Slovenia	This site is known worldwide for the earliest fossil records of seahorses.^[194]	Two fossil seahorse species, Hippocampus sarmaticus and H. slovenicus from this site.
Ashfall Fossil Beds	11.83 Ma	Nebraska, US	The Ashfall Fossil Beds of Antelope County in northeastern Nebraska are rare fossil sites of the type called lagerstätten that, due to extraordinary local conditions, capture an ecological "snapshot" in time of a range of well-preserved fossilized organisms. Ash from a Yellowstone hotspot eruption 10-12 million years ago created these fossilized bone beds.	A bone-bed containing the fossils of the basal rhino Teleoceras and the three-toed horse Cormohipparion.
Alcoota Fossil Beds	8 Ma	Northern Territory, Australia	It is notable for the occurrence of well-preserved, rare, Miocene vertebrate fossils, which provide evidence of the evolution of the Northern Territory's fauna and climate. The Alcoota Fossil Beds are also significant as a research and teaching site for palaeontology students.	Miscellaneous fossils of several macropod marsupials
Saint-Bauzile	7.6-7.2 Ma	Ardèche, France	A Late Miocene site preserving articulated mammal skeletons with skin and fur impressions.^[195]
Capo San Marco Formation	~7 Ma	Sardinia, Italy	A microbialite containing exceptionally preserved Girvanella-type filaments.^[196]
Tresjuncos	6 Ma	Cuenca, Spain	A Late Miocene lacustrine konservat-lagerstätte containing fossils of diatoms, plants, crustaceans, insects, and amphibians.^[197]
Gray Fossil Site	4.9-4.5 Ma	Tennessee, US	As the first site of its age known from the Appalachian region, the Gray Fossil Site is a unique window into the past. Research at the site has yielded many surprising discoveries, including new species of red panda, rhinoceros, pond turtle, hickory tree, and more. The site also hosts the world's largest known assemblage of fossil tapirs.	Fossil skull of Pristinailurus, a North American relative of the modern Red Panda
Willershausen	Late Pliocene	Lower Saxony, Germany	A lacustrine fossil site containing well preserved beetles.^[198]

Quaternary

Site(s)	Age	Location	Significance	Notable fossils/organisms
The Mammoth Site	26 Ka	South Dakota, US	The facility encloses a prehistoric sinkhole that formed and was slowly filled with sediments during the Pleistocene era. As of 2016, the remains of 61 mammoths, including 58 North American Columbian and 3 woolly mammoths had been recovered. Mammoth bones were found at the site in 1974, and a museum and building enclosing the site were established.	Fossil skeleton of Arctodus simus, a large species of "short-faced" bear that was one of North America's largest predators during the Pleistocene.
Rancho La Brea Tar Pits	40–12 Ka	California, US	A group of tar pits where natural asphalt (also called asphaltum, bitumen, or pitch; brea in Spanish) has seeped up from the ground for tens of thousands of years. Over many centuries, the bones of trapped animals have been preserved. Among the prehistoric species associated with the La Brea Tar Pits are Pleistocene mammoths, dire wolves, short-faced bears, American lions, ground sloths, and, the state fossil of California, the saber-toothed cat (Smilodon fatalis).	Fossil skeleton of Mammuthus columbi excavated from the tar pits
Waco Mammoth National Monument	65–51 Ka	Texas, US	A paleontological site and museum in Waco, Texas, United States where fossils of 24 Columbian mammoths (Mammuthus columbi) and other mammals from the Pleistocene Epoch have been uncovered. The site is the largest known concentration of mammoths dying from a (possibly) reoccurring event, which is believed to have been a flash flood.	Fossil skeleton of a mammoth found at the Waco site
El Breal de Orocual	2.5–1 Ma	Monagas, Venezuela	The largest asphalt well on the planet. Like the La Brea Tar Pits, this site preserves a number of megafauna like toxodonts, glyptodonts, camelids, and the felid Homotherium venezuelensis.	Restoration of the environment at El Breal de Orocual, showing Glyptodon, Coragyps, Dasypus and Myrmecophaga
El Mene de Inciarte	28–25.5 Ka	Zulia, Venezuela	Another series of tar pits. These also preserve a similar assemblage of megafauna.
Naracoorte Caves	500-1 Ka	South Australia, Australia	A series of caves that preserve numerous pleistocene megafauna, like Thylacoleo, and is recognized as a World heritage site alongside the older, but geographically similar Riversleigh site.	Skeleton of Thylacoleo at the Naracoorte Caves
Mare aux Songes	4 Ka	Mauritius	A marsh that preserves a diversity of subfossil animals and plants, many of which were driven to extinction without proper documentation following human arrival, most notably the famous dodo. The mortality assemblages may have formed from a freshwater lake that was occasionally impacted by catastrophic droughts.^[199]	Dodo skeleton from Mare aux Songes
Crawford Lake	800 Ya-present	Ontario, Canada	This lake is notable for its detailed preservation of rotifer and dinoflagellate fossils even after centuries, documenting the ecological shifts that occurred in and around the lake following the establishment of a Iroquoian village from 1268–1486 CE, and later following European colonization of the region in the early 19th century.^[200]