AFossil-Lagerstätte (German:[ˈlaːɡɐˌʃtɛtə], fromLager 'storage, lair'Stätte 'place'; pluralLagerstätten) is asedimentary deposit that exhibits extraordinaryfossils with exceptional preservation—sometimes including preserved soft tissues. These formations may have resulted from carcass burial in ananoxic 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 spansgeological time from theNeoproterozoic era to thepresent.
Worldwide, some of the best examples of near-perfect fossilization are theCambrianMaotianshan shales andBurgess Shale, theOrdovicianSoom Shale, theSilurianWaukesha Biota, theDevonianHunsrück Slates andGogo Formation, theCarboniferousMazon Creek, the TriassicMadygen Formation, theJurassicPosidonia Shale andSolnhofen Limestone, theCretaceousYixian,Santana, &Agua Nueva formations and theTanis Fossil Site, theEoceneFur Formation,Green River Formation,Messel Formation &Monte Bolca, theMioceneFoulden Maar andAshfall Fossil Beds, thePlioceneGray Fossil Site, and thePleistoceneNaracoorte Caves &La Brea Tar Pits.
Palaeontologists distinguish two kinds:[1][2]
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Konservat-Lagerstätten preserve lightlysclerotized 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 thepalaeoecology of ancient aquatic communities. In 1986,Simon Conway Morris calculated only about 14% of genera in the Burgess Shale had possessedbiomineralized tissues in life. The affinities of the shelly elements ofconodonts were mysterious until the associated soft tissues were discovered near Edinburgh, Scotland, in the Granton LowerOil Shale of theCarboniferous.[3] Information from the broader range of organisms found inLagerstätten have contributed to recentphylogenetic reconstructions of some majormetazoan groups.Lagerstätten seem to be temporally autocorrelated, perhaps because global environmental factors such as climate might affect their deposition.[4]
A number oftaphonomic pathways may produceKonservat-Lagerstätten:[5]
The identification of a fossil site as aKonservat-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 sedimentfacies encouraging preservation.[5]
The world's major Lagerstätten include:
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Francevillian B Formation | 2140-2080 Ma | Franceville, Gabon | Possibly preserves the earliest macroscopic eukaryotes,[6][7] however these may instead be pseudofossils.[8][9] |  |
| Chuanlinggou Formation | 1630 Ma[10] | Yan Mountains, North China | Preserves the first definitive multicellular eukaryotes in the form ofQingshania,[10]Tawuia andChangchengia.[11] |  |
| Chitrakoot Formation | 1600 Ma | Vindhya Range, India | Preserves earliest red algae,Rafatazmia andRamathallus[12] |  |
| Gaoyuzhuang Formation | 1588.8±6.5 Ma[13] | Yan Mountains, North China | Preserves some of the first algae such asTuanshanzia andGrandilingulata,[14] alongside a diverse microbiota including possible fungi.[15] |  |
| Volyn biota | 1500 Ma | Zhytomyr Oblast, Ukraine | Possibly one of the first fossil sites of thedeep biosphere.[16][17] |  |
| Hunting Formation | 1200 Ma | Somerset Island, Canada | Shallow-water Mesoproterozoic deposits, containing multiple genera of algae[18][19] includingBangiomorpha. | .png) |
| Nonesuch Formation | 1083-1070 Ma | Michigan, USA | An oxygenated Mesoproterozoic lake[20] containing exceptionally preserved limnic microbes.[21] | |
Lakhanda Lagerstätte | 1030-1000 Ma | Uchur-Maya Depression, Russia | A site preserving a Mesoproterozoic community dominated by anaerobic bacteria.[22] The lagerstätte contains evidence of trophic interactions from theBoring Billion.[23][24] | |
1000–850Ma | South Australia | Preserved fossils includecyanobacteriamicrofossils. | _(Neoproterozoic%2c_%7e800_Ma%3b_Bitter_Springs%2c_Australia)_1_(41559714260).jpg) | |
| Diabaig Formation | 994 ± 48 Ma[25] | Scotland | A freshwater environment preserving phosphatic microfossils,[26] which represent some of the oldest known non-marine eukaryotes.[27] | |
| Dolores Creek Formation | 950 Ma | Yukon, Canada | An Early Tonian site containing pyritised macroalgal fossils.[28] | |
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.[29] | |
600–555 Ma | Guizhou Province, China | Spans the poorly understood interval between the end of theCryogenianperiod and the late EdiacaranAvalon explosion. |  | |
| Portfjeld Formation | 570 Ma | North Greenland | A Middle Ediacaran biota from the continent ofLaurentia exhibiting Doushantuo-type preservation.[30] | |
565 Ma | Newfoundland, Canada | This site contains one of the most diverse and well-preserved collections ofPrecambrianfossils. |  | |
| Itajai Biota | 563 Ma | Brazil | An Ediacaran lagerstatte preserved by volcanism.[31] | |
555 Ma | South Australia | The type location theEdiacaran period, and has preserved a significant amount of fossils from that time. |  | |
| Shibantan Lagerstätte | 551-543 Ma | Hubei, China | A terminal Ediacaran fossil assemblage preserving life forms living just before the Proterozoic-Phanerozoic transition.[32] | |
| Gaojiashan Lagerstätte | 551-541 Ma | Shaanxi, China | A lagerstätte documenting tube growth patterns ofCloudina.[33] | |
| Jiucheng Member | 551-543 Ma | Yunnan, China | A latest Ediacaran macrofossil biota dominated by giant, unbranching thallophytes.[34] | |
Khatyspyt Lagerstätte | 544 Ma | Yakutia, Russia | A Late Ediacaran lagerstätte preserving an Avalon-type biota.[35] | |
| Bernashivka open pit | ? (UpperVendian) | Vinnytsia Oblast, Ukraine | A Late Ediacaran lagerstätte with numerous soft-bodied animals, algae, microfossils, bacteria, and fungi, comprising a number of different geological formations.[36] |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Zhangjiagou | Fortunian | Shaanxi, China | A lagerstätte from the earliest Cambrian notable for its fossils of cnidarians,[37] cycloneuralians,[38][39] and the basal ecdysozoanSaccorhytus coronarius.[40] |  |
Maotianshan Shales (Chengjiang) | 518 Ma | Yunnan, China | The preservation of an extremely diverse faunal assemblage renders the Maotianshan Shales the world's most important formation for understanding the evolution of earlymulti-cellular life. Microscopic animals likeYicaris are preserved here, showing the presence of an Orsten-type deposit within the formation.[41] |  |
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. |  | |
523-518 Ma | A site known for its fauna, and that they were most likely preserved by adeath mask. It is a part of the largerBuen Formation, and has a fauna similar to the Maotianshan shales. |  | ||
| Xiaoshiba Lagerstätte | Cambrian Stage 3 | Yunnan, China | A site known for its detailed preservation of Early Cambrian macroalgae,[42] alongside fossils ofOmnidens.[43] |  |
| 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 formerLaurentia. 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.[44] |  |
518 Ma | One of the oldest known Cambrianlagerstätten. The fauna of this site is unique, as it seems that they were adapted to living in dysaerobic conditions.[45] |  | ||
EasternYunnan, China | A relatively diverse lagerstätte within theWulongqing Formation, between the more famous Maotianshan Shales and Burgess Shale in age, alongside having both taxa from the previous two formations and completely new genera/species preserved here. It is also unusual for being quite shallow and having a brachiopod-dominated fauna.[46] |  | ||
| Cranbrook Lagerstätte | Cambrian Stage 4 | British Columbia | One of the oldest Burgess Shale-type biotas of North America.[47] | |
| 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.[48] |  |
513 Ma | Noted soft tissue mineralization, most often of blockyapatite or fibrouscalcium carbonate, including the oldest phosphatized muscle tissue. |  | ||
| 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.[49] | |
513–501 Ma | Guizhou, China | The middle part of the Kaili Formation, theOryctocephalus indicus Zone, contains a Burgess Shale-type lagerstätte with many well-preserved fossils known collectively as the Kaili Biota. | _(Kaili_Formation%2c_Middle_Cambrian%3b_quarry_between_Balang_%2526_Dunzhou%2c_Guizhou_Province%2c_China).jpg) | |
Murero Lagerstätte | 511-503 Ma | Spain | Thanks to the paleontological content, mainly trilobites, fourteenbiozones 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.[50] |  |
~510–500 Ma | CentralWisconsin, 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 largemollusks andeuthycarcinoid arthropods). Other notable fossils include strandedscyphozoans, and some of the oldest true crustaceans (in the form ofphyllocarids). |  | |
| Henson Gletscher Formation | Wuliuan | North Greenland | A phosphatised lagerstätte preserving hatching priapulid larvae, pentastomids[51] and abundantbradoriid andphosphatocopid arthropods.[52] | |
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. |  | |
~505 Ma? | SoutheasternBritish Columbia | Similar to the Burgess Shale above (both formations even have similar depositional environments next to escarpments), but seemingly lower-diversity and slightly younger. Several organisms are shared between both formations, although the Duchesnay Formation preserves several new genera. Brine pools are also preserved.[53] | ||
507 Ma | 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. |  | ||
Linyi Lagerstätte | 504 Ma | A lagerstätte recognised for its exceptional preservation of arthropod limbs, intestines, and eyes.[54] |  | |
| Ravens Throat River Lagerstätte | Drumian | Northwest Territories | A Burgess Shale-type biota coeval in age with the more famous Wheeler Shale and Marjum Formation.[55] | |
504 Ma | WesternUtah, US | A world-famous locality known for its prolificagnostid andElrathia kingii trilobite remains. Varied soft bodied organisms are also locally preserved, includingNaraoia,Wiwaxia andHallucigenia. |  | |
502 Ma | WesternUtah, US | A site known for its occasional preservation of soft-bodied tissue, and diverse assemblage. |  | |
500 Ma | WesternUtah, US | A site that is dominated by trilobites andbrachiopods, but also comprising various soft-bodied organisms, such asFalcatamacaris. |  | |
500 Ma | The Orsten sites reveals the oldest well-documentedbenthicmeiofauna in the fossil record. Fossils such as microfossils of arthropods like free-livingpentastomids are known. Multiple "Orsten-type"lagerstätten are also known from other countries. |  |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
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 CambrianBurgess shale-type deposits. |  | |
| 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.[56] | |
Liexi fauna | About 470 Ma (early-middleFloian) | Hunan Province, China | Preserves Early Ordovician fauna with soft tissue, includes not only Cambrian relics but also taxa originated during Ordovician.[57] | |
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. |  | |
460 Ma | Tennessee, US | Low-diversity assemblage of arthropod fossils, which are preserved well because of volcanic ash. |  | |
460 Ma | 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.[59] |  | ||
460? Ma | New York, US | Noted exceptionally preserved trilobites with soft tissue preserved bypyrite replacement. |  | |
? (Sandbian) | Colorado, US | Although preservation is not excellent, thislagoonal site provides early vertebrate fossils such asAstraspis andEriptychius. |  | |
about 455? Ma | New York, US | This site is an excellent example of an obrution (rapid burial or "smothered")Lagerstätte. | _(16763508314).jpg) | |
| 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). |  |
Brechin Lagerstätte | 450 Ma | Ontario, Canada | Known for preserving one of the most diverse crinoid fauna of theKatian.[60] |  |
450? Ma | South Africa | Known for its remarkable preservation of soft-tissue in fossil material. Deposited in still waters, the unit lacksbioturbation, perhaps indicatinganoxic conditions. |  | |
Tafilalt, Morocco | Known from range of non-biomineralised and soft-bodied organisms in polar environment.[61] |  | ||
? (middleKatian) | Manitoba, Canada | Fossils like algae,conulariids and trilobites are known from this site. | ||
449-445.6 Ma | Manitoulin District, Canada | Low-diversity assemblage of arthropod fossils. |  | |
William Lake (Stony Mountain Formation)[58] | 445 Ma | Manitoba, Canada | Well-preserved fossils likejellyfish,xiphosurans,sea spiders are known from this site, it is important since many of the fossils are unknown in other Ordovician sites. |  |
Airport Cove[58] | 445 Ma | Manitoba, Canada | Fossils like eurypterids, algae and xiphosurans are preserved in this site. |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
Kalana Lagerstätte | ~440 Ma | Estonia | Known for well preserved fossils of algae and crinoids,[62] along with an osteostracan fossilised via an extremely unusual carbonaceous mode of preservation that was previously unknown among vertebrates.[63] |  |
Chongqing Lagerstätte (Huixingshao Formation)[64] | 436 Ma | Chongqing, China | This site preserved complete fossils of earliestjawed vertebrates, as well as somegaleaspids andeurypterids. |  |
~435 Ma | Wisconsin, US | Well-studied site known for the exceptional preservation of its diverse, soft-bodied and lightly skeletonized fauna, includes many majortaxa found nowhere else in strata of similar age. It was one of the first fossil sites with soft bodied preservation known to science. |  | |
Herefordshire Lagerstätte (Coalbrookdale Formation) | ~430 Ma | Known for the well-preserved fossils of variousinvertebrate 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 toPompeii.[65] Some of the fossils are regarded as earliest evidences and evolutionary origin of some of the major groups of modern animals. |  | |
~425 Ma | Known for preservation of both hard and soft bodied organisms in great detail, including early scorpions, eurypterids, agnathan vertebrates, and several other species. |  | ||
422.9-416 Ma | Ontario & New York State | This limestone have produced thousands of fossileurypterids, such as giantAcutiramus and well-knownEurypterus, as well as other fauna like scorpions and fish. |  | |
~420 Ma | Pennsylvania, US | Known from exceptionally preserved mass assemblage ofEurypterus, the most abundant eurypterid in the fossil record. |  | |
415 Ma[67] | New York, US andOntario, Canada | Echinoderms (such as crinoids) and trilobites are known from Lewiston Member in this shale. | _(15273506362).jpg) |
| 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 overlyingvolcanic deposit and hot springs. As well as one of the first known fully terrestrial ecosystems. |  |
| Waxweiler Lagerstätte (Klerf Formation) | 409-392 Ma | Eifel,Germany | Waxweiler Lagerstätte is known from well-preserved fossils ofchelicerates, giant claw ofJaekelopterus rhenaniae shows the largest arthropod ever known. |  |
| Heckelmann Mill | 395 Ma | Rhineland-Palatinate, Germany | Heckelmann Mill preserves well preserved rhinocaridid archaeostracan phyllocarids,[68] along with exceptionally abundant crinoid holdfasts from the lateEmsian.[69] | |
| Hunsrück Slates (Bundenbach) | 390 Ma | Rheinland-Pfalz, Germany | The Hunsrück slates are one of the few marine Devonianlagerstätte having soft tissue preservation, and in many cases fossils are coated by a pyritic surface layer. |  |
| 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 offish have been described from the formation, and arthropods. |  |
| 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 earlytetrapods. |  |
| Kowala Lagerstätte | ~368 Ma | Świętokrzyskie Voivodeship,Poland | A Late Devonian site known for its fossils of non-biomineralised algae and arthropods.[70] | |
| Maïder Basin | 368 Ma (for Thylacocephalan Layer) | Anti-Atlas,Morocco | Thylacocephalan Layer and Hangenberg Black Shale in this basin provides well-preserved fossils ofFamennian fauna, includingchondrichthyans andplacoderms that preserved soft tissues.[71] |  |
| Strud[72] | ? (LateFamennian) | Namur Province,Belgium | Mainly juvenile placoderms are known, suggesting this site would be nursery site of placoderms.[73] Various biota like tetrapods, arthropods and plants are also known,Strudiella from this site may be the earliest insect, but its affinity is disputed. |  |
| Canowindra, New South Wales (Mandagery Sandstone) | 360 Ma | Australia | An accidentally discoveredlagerstätte known for its exceptional preservation ofSarcopterygian andPlacoderm fish. |  |
| Waterloo Farm Lagerstätte (Witpoort Formation) | 360 Ma | South Africa | Important site that providing the only record of a high latitude (nearpolar) coastal ecosystem, overturning numerous assumptions about high latitude conditions during the latest Devonian. |  |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Granton Shrimp Bed | ? (Dinantian) | Firth of Forth, Scotland | Dominated by well-preservedcrustacean fossils, this site provided first body fossil ofClydagnathus which solved long-lasted mystery ofconodont fossils. | |
| East Kirkton Quarry[74] | 335 Ma | West Lothian, Scotland | This site has produced numerous well-preserved fossils of early tetrapods liketemnospondyls orreptiliomorphs, and large arthropods like scorpions or eurypterids. |  |
| Bear Gulch Limestone | 324 Ma | Montana, US | Alimestone-richgeological lens in centralMontana. It is renowned for its unusual and ecologically diverse fossil composition ofchondrichthyans, the group of cartilaginous fish containing modernsharks,rays, andchimaeras. Other animals like brachiopods,ray finned fish, arthropods, and the possible molluskTyphloesus are also known from the site. |  |
| Wamsutta Formation | ~320-318 Ma | Massachusetts | A subhumid alluvian fan deposit that preserves ichnofossils, plants, invertebrates, and vertebrates.[75] | |
| Bickershaw[76] | ? (Langsettian) | Lancashire, England | This locality contains exceptionally preserved fossils withinnodules. Arthropods have greater diversity, many of which are aquatic ones that lived in brackish environment. |  |
| 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 knownsauropsid,Hylonomus. |  |
| Castlecomer fauna | 315-307 | Kilkenny,Ireland | Akonservat-lagerstätte with a high number of well-preservedspinicaudatan clam shrimp.[77][78] | |
| Linton Diamond Coal Mine[79][80] | 310 Ma | Ohio, US | A site known for its number of prehistorictetrapods, like thelepospondylDiceratosaurus.[81] |  |
| Mazon Creek | 310 Ma | Illinois, US | A conservationlagerstätte found nearMorris, inGrundy County, Illinois. The fossils from this site are preserved inironstoneconcretions 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 animalTullimonstrum, is only known from these deposits. |  |
| Buckhorn Asphalt Quarry | ~310 Ma | Oklahoma, USA | A quarry of theBoggy Formation known for its exceptionally rich orthocerid assemblage.[82] | |
| 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 andhybodonts, toactinopterygians andsarcopterygians.[83] |  |
| Campáleo Outcrop | Gzhelian | Santa Catarina, Brazil | A fungal and palynological lagerstätte from Gondwana duringLate Palaeozoic Ice Age.[84] | |
| Montceau-les-Mines | 300 Ma | France | Exceptional preservation ofLate Carboniferous fossil biota are known, including various vertebrates and arthropods, as well as plants.[85][86] |  |
| 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. |  |
| Carrizo Arroyo | ? (LatestGzhelian to earliestAsselian) | New Mexico, US | This site is known from exceptional preservation of arthropod fossils, mainly insects.[87] |
| 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.[88][89] | |
| Meisenheim Formation[90] | ?(Asselian to earlySakmarian)[91] | Lebach, Germany | This site is well-known for the rich occurrence of fauna lived in large freshwater lakes, including fish,temnospondyls, insects and others. |  |
| Franchesse | 292 Ma | Massif Central, France | ASakmarianseymouriamorph lagerstätte from the Bourbon l'Archambault Basin in the French Massif Central containing hundreds of complete seymouriamorph specimens.[92] | |
| Chemnitz petrified forest | 291 Ma | Saxony, Germany | Apetrified forest in Germany that is composed ofArthropitys bistriata, a type ofCalamites, gianthorsetails that are ancestors of modernhorsetails, found on this location with never seen multiple branches. Many more plants and animals from this excavation are still in an ongoing research.[93] |  |
| Mangrullo Formation | about 285–275 Ma (Artinskian) | Uruguay | This site is known for its abundantmesosaur fossils. It also contains the oldest knownkonservat-lagerstätte inSouth America, as well as the oldest known fossils ofamnioteembryos.[94] |  |
| 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.[95] |  |
| 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.[96] | |
| Onder Karoo | 266.9–264.28 Ma | Karoo Basin, South Africa | A high latitude, cool-temperate lacustrine ecosystem preserving detailed plant and insect fossils.[97] | |
| Sakamena Group[98] | 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 glidingneodiapsids. |  |
| 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. |  |
| Huopu Lagerstätte | ~255 Ma | Guizhou, China | A plant fossil site documenting floral dynamics between theend-Guadalupian andend-Permian extinction events.[99] |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Guiyang biota[100] | 250.8 Ma | Guizhou Province, China | The oldest knownMesozoic lagerstätte (Dienerian). It preservestaxa belonging to 12classes and 19orders, including several species of fish. |  |
| Paris biota[101] | ~249 Ma | Idaho,Nevada, USA | This earliestSpathian aged assemblage preserves fossils belonging to 7phyla and 20orders, combiningPaleozoic groups (e.g. leptomitid protomonaxonidsponges otherwise known from the early Paleozoic) with members of the Modern evolutionary fauna (e.g. gladius-bearingcoleoids). |  |
| Jialingjiang Formation[102] | 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 likefilter feeding. |  |
| Nanlinghu Formation[102] | 248 Ma | Anhui Province, China | This site provides important fossils to show early evolution ofichthyosauriforms. |  |
| Petropavlovka Formation | 248 Ma | Orenburg Oblast, Russia | A site known for preservingoligochaetes, whose fossil record is extremely sparse.[103] | |
| Zarzaïtine Formation | Olenekian-Anisian | In Amenas,Algeria | A site with a high number of exceptionally well-preservedtemnospondyl specimens, indicating of a seasonal climate with sudden droughts, with a freshwater ecosystem that could rapidly turn into asebkha.[104] | |
| Luoping Biota (Guanling Formation)[105] | ~247-245 Ma | Yunnan, China | Various marine animals are preserved in this site, showing how marine ecosystem recovered after Permian extinction.[106] |  |
| Hawkesbury Sandstone[107] | Anisian | Sydney, Australia | This sandstone produced many of freshwater fish fossils, as well as arthropods. |  |
| Grès à Voltzia | 245 Ma | France | A fossil site remarkable for its detailed myriapod specimens.[108] It also contains the earliest knownaphid fossils.[109] | |
| 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 byichthyosaurs.[106] |  |
| Vossenveld Formation | ? (Anisian) | Winterswijk,Netherlands | An exposure of thisMuschelkalk 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.[106] | |
| Strelovec Formation | ? (Anisian) | Slovenia | A formation with well-preserved Triassichorseshoe crabs.[110] | |
| Saharonim Formation | Late Anisian/Lower Ladinian | Southern District,Israel | Onebrachiopod-dominated horizon of this formation documents the rapid burial of a community of exclusively juvenileCoenothyris brachiopods and ten different bivalve genera.[111] | |
| Besano Formation[102] | 242 Ma | Alps,Italy andSwitzerland | This formation is designated as aWorld Heritage Site, as it is famous for its preservation of Middle Triassic marine life including fish and aquatic reptiles.[106][112] |  |
| Pelsa-Vazzoler Lagerstätte | Late Ladinian | Northeastern Italy | A site preserving a megadiverse mollusc fauna during the recovery from the Permian-Triassic extinction event.[113] | |
| Xingyi biota (Zhuganpo Formation)[102] | ? (UpperLadinian - LowerCarnian) | Guizhou andYunnan, China | Previously considered as part of Falang Formation, this site yields many articulated skeletons of marine reptiles, as well as fish and invertebrates. |  |
| Guanling biota (Xiaowa Formation)[102] | ? (Carnian) | Guizhou, China | Like Xingi Biota, this site also yields well-preserved marine fauna, especially many species ofthalattosaurs are known. |  |
| Polzberg | 233 Ma | Austria | A site from theReingrabener Schiefer known for exceptional preservation of bromalites[114] and of cartilage,[115] deposited during theCarnian Pluvial Event.[116] |  |
| 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 andsynapsids have been recovered from the formation too, as well as minor fossil flora. |  |
| Cow Branch Formation | 230 Ma | Virginia, US | This site preserves a wide variety of organisms (including Fish, reptiles,arachnids, andinsects). |  |
| Alakir Çay | ? (Norian) | southwestTurkey | Akonservat-lagerstätte with exceptionally well-preserved Triassic corals, retaining much of their original aragonite skeletons.[117] |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Yuzhou Biota (Ziliujing Formation) | ~199 Ma | Chongqing, China | A fossil biota representing an extremely well-preserved lake ecosystem with freshwater mollusks, ray-finned fishes, lungfishes, sharks, and evenpliosauroids, documenting one of the earliest well-preserved terrestrial ecosystems from Asia following theTriassic-Jurassic extinction event.[118] |  |
| Osteno (Moltrasio Formation)[119] | ~196-188 Ma | Italy | Several kinds of marine biota such as fish, crustaceans, cephalopods, polychaetes, andnematodes have been recovered. This site is the only fossil deposit in Italy in which soft tissues are preserved other thanMonte Bolca. |  |
| Charmouth Mudstone Formation | ~196-188 Ma | Lyme Regis andCharmouth (UK) | Know for its amniote remains, specially complete specimens of the armoured dinosaurScelidosaurus, but alsoplesiosaurs &ichthyosaurs.[120][121][122] Large quantities of exceptional fish specimens and a diverse entomofauna are also known.[123] | .jpg) |
| Rotzo Formation | ~192-184 Ma | Italy | Diverse fossils from a Carbonate archipelago, specially withinplicatostylid reefs. It is mostly known for the exceptional preservation of organisms likeTestate amoebae, and specially due to itsamber-bearing layers and associated macroflora.[124][125] | |
| Calcaires du Bou Dahar | 196-183 Ma | Morocco | It records the evolution of a carbonate platform.[126] It is known mostly for its rich coral bioherms and associated invertebrates.[127][128] | |
| 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,[129] and for being deposited during theToarcian Oceanic Anoxic Event (TOAE).[130][131][132] | |
| Strawberry Bank | 183 Ma | Somerset, England | A site from the TOAE documenting marine life during the recovery from theTriassic-Jurassic extinction event as well as the turmoil of the TOAE.[133] The oldest pseudoplanktonic barnacles in the fossil record,[134] near-complete ichthyosaur skeletons,[135] and evidence of ichthyosaur niche partitioning are preserved at this site.[136] | |
| Grimmen (Ciechocinek Formation) | 183 Ma | Grimmen,Germany | A coastal marine sequence, know specially for its fish fauna.[137][138] The formation is also known for its rich entomofauna, including collections of up to 3000 specimens.[139] | |
| 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.[140][141] |  |
| Cabeço da Ladeira | Late Bajocian | Portugal | A site known for exquisite preservation of microbial mats in a tidal flat.[142] | |
| Monte Fallano | ? (Bajocian-Bathonian) | Campania, Italy | ThisPlattenkalk preserves fossils of terrestrial plants, crustaceans and fish.[143] | |
| Christian Malford | Callovian | Wiltshire, England | A site in the Oxford Clay Formation which preserves exceptionally detailed coleoid fossils.[144] | |
| Mesa Chelonia[145] | 164.6 Ma | Shanshan County, China | This site is notable because it contains a large turtlebonebed, containing specimens of the genusAnnemys. 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 theArdèche region of southwesternFrance, offers paleontologists an outstanding view of an undisturbedpaleoecosystem that was preserved in fine detail. Notable finds includeretinal structures in the eyes ofthylacocephalan arthropods, and fossilized relatives of the modern dayvampire squid, likeVampyronassa rhodanica. |  |
| Shar Teeg Beds | 160-145 Mya | Govi-Altay,Mongolia | Many insect remains and some vertebrates like relatives of crocodilians are known from this site.[146] | |
| 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, includingpterosaurs,salamanders,lizards andcrocodiles. | .JPG) |
| 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 ofpyroclastic rock interspersed with basic volcanic andsedimentary rocks. |  |
| La Casita Formation | Kimmeridgian | Coahuila, Mexico | A marinekonzentrat-lagerstätte deposited in a hemipelagic mud bottom during dysoxic conditions.[147] | |
| Talbragar fossil site[148] | 151 Ma | New South Wales, Australia | This bed is part ofPurlawaugh Formation, and provided fauna like fish and insects that lived around the lake. |  |
| 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 ofJurassic dinosaur fossils ever found, is a paleontological site located nearCleveland, Utah, in theSan Rafael Swell, a part of thegeological layers known as theMorrison Formation. Up to 15,000 have been excavated from this site alone. |  |
| Canjuers Lagerstätte | 150 Ma | France | This site shows a high amount of biodiversity, including reptiles, invertebrates, fish, and other organisms. |  |
| Agardhfjellet Formation | 150-140 Ma | Spitsbergen, Norway | The formation contains the Slottsmøya Member, a highly fossiliferous unit where manyichthyosaur andplesiosaur fossils have been found, as well as abundant and well preserved fossils of invertebrates. |  |
| Solnhofen Limestone | 149-148 Ma | Bavaria, Germany | This site is unique as it preserves a rare assemblage offossilized organisms, including highly detailed imprints of soft bodied organisms such assea jellies. The most familiar fossils of theSolnhofen Plattenkalk include the early featheredtheropod dinosaurArchaeopteryx preserved in such detail that they are among the most famous and most beautiful fossils in the world. | |
| Owadów–Brzezinki site | ~148 Ma | Łódź Voivodeship, Poland | A marine deposit of theKcynia Formation similar to the Solnhofen Formation, with large numbers of preserved insect remains, numerous marine invertebrates, and vertebrates including fishes, marine reptiles, and pterosaurs.[149][150] |  |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Muzinho Shale | latest Jurassic/earliest Cretaceous | Parnaíba Basin, Brazil | A black shale deposit containing articulated, three-dimensionally preserved fish skeletons.[151] | |
| Angeac-Charente bonebed | ~141 Ma | Charente, France | A lagerstätte preserving both vertebrate and invertebrate fossils from the poorly representedBerriasian stage known for its taphonomic and sedimentological ‘frozen scenes’.[152] | |
| 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.[153] |  |
| Lebanese amber | ~130-125 Ma (Barremian) | Lebanon | Preserves a high diversity of insects from theEarly Cretaceous, and is among the oldest known fossilized amber to contain a significant number of preserved organisms.[154] Includes many of the oldest known members of modern insect groups, and many of the youngest known members for extinct insect groups.[155] |  |
| Lebanese dysodiles | ~125 Ma (early Barremian) | Lebanon | A deposit of dysodiles (sedimentary rocks with high organic matter content) representing a well-preserved freshwater ecosystem, including plants, pollen grains, invertebrates (including insects), turtles, and a diverse fish fauna.[156] | |
| Las Hoyas | about 125 Ma (Barremian) | Cuenca, Spain | The site is mostly known for its exquisitely preserved dinosaurs, especiallyenantiornithines. The lithology of the formation mostly consists oflacustarinelimestone deposited in a freshwater wetland environment. |  |
| 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 tyrannosauroidYutyrannus, the therizinosaurBeipiaosaurus, and various small birds, along with a selection of other dinosaurs, such as the iguanodontianBolong, the sauropodDongbeititan and the ceratopsianPsittacosaurus. Other biota included the troodontidMei, the dromaeosauridTianyuraptor, and the compsognathidSinosauropteryx. |  |
| 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 ceratopsianPsittacosaurus and the early birdConfuciusornis, both of which are also found in the Yixian Formation. Also notable are the very abundant specimens of the dromaeosauridMicroraptor, which is known from up to 300 specimens and is among the most common animals found here. |  |
| 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.[157] | |
| 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. |  |
| Xiagou Formation | about 120–115? Ma | Gansu, China | This site is known outside the specialized world of Chinesegeology as the site of a lagerstätte in which the fossils were preserved ofGansus yumenensis, the earliest true modern bird. |  |
| Paja Formation | 130-113 Ma | Colombia | This site is famous for its vertebrate fossils and is the richest Mesozoicfossiliferous 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 sauropodPadillasaurus. |  |
| Koonwarra Fossil Bed[158] | around 118-115 Ma | Victoria, Australia | This site is composed ofmudstone sediment thought to have been laid down in a freshwater lake. Arthropods, fish and plant fossils are known from this site. |  |
| Crato Formation | 113 Ma | northeast Brazil | The Crato Formation earns the designation of lagerstätte due to an exceedingly well preserved and diversefossil 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 ofpterosaurs, reptiles and amphibians, invertebrates (particularly insects), and plants. Also known from this site isUbirajara, the first non-avian dinosaur from the southern hemisphere with evidence of feathers. Additionally, the formation abounds with evidence of plant-insect interaction.[159] |  |
| Amargosa Bed (Marizal Formation) | ? (Aptian-Albian) | northeast Brazil | Fluvial site which preserved fish, crustacean and plant fossils.[160] | |
| Pietraroja Plattenkalk | 113-110 Ma | Campania, Italy | Akonservat-lagerstätte famous for its diverse and well-preserved fish and plant fossils. Also known from this formation isScipionyx, one of Europe's most well-preserved dinosaurs.[161][162] |  |
| Jinju Formation | 112.4–106.5 Ma | South Korea | The Jinju Formation is notable for the post-Jehol Group insect assemblage,[163] as well as other fauna such asisopods and fish.[164][165] The site is also notable for its abundance and diversity oftetrapod trackways.[166] |  |
| Tlayúa Formation | 110 Ma | Puebla,Mexico | A marine lagerstätte preservingAlbian actinopterygians and lepidosaurs.[167] |  |
| Romualdo Formation | 108–92 Ma | northeast Brazil | The Romualdo Formation is a part of theSantana Group and has provided a rich assemblage of fossils; flora, fish, arthropods insects, turtles, snakes, dinosaurs, such asIrritator, and pterosaurs, including the genusThalassodromeus. The stratigraphic units of the group contained several feathers of birds, among those the first record ofMesozoic birds in Brazil. |  |
| Muhi Quarry (El Doctor Formation) | ? (Albian toCenomanian, probably Late Albian)[168] | 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. |  |
| Puy-Puy Lagerstätte | 100.5 Ma | France | A paralic site preserving a variety of ichnofossils,[169] along with some vertebrate remains.[170] The site preserves evidence of plant-insect interaction.[171] | |
| Burmese amber | 101-99 Ma (latest Albian/earliestCenomanian) | Myanmar | More than 1,000 species of taxa have been described from ambers fromHukawng 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. |  |
| English Chalk | 100-90 Ma (Cenomanian toTuronian) | 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.[172] | .jpg) |
| Haqel/Hjoula/al-Nammoura | 95-94 Ma | Lebanon | Famous Lebanesekonservat-lagerstätten of the LateCretaceous (middle to lateCenomanian) 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 ofoctopuses.[173] |  |
| Ein Yabrud (Amminadav Formation orBet-Meir Formation) | Cenomanian | Palestine | The marine fossil site with well-preserved marine vertebrate fossils, especially fossils of early legged snakes likeHaasiophis andPachyrhachis has quality of preservation rivals that of other famous Lagerstatten.[174] |  |
| Komen Limestone | 95-94 Ma | Komen,Slovenia | A Late Cenomanian locality in theKarst of Slovenia with a high diversity of articulated fossil fish, in addition to small reptiles and invertebrates.[175] | _(16142481744).jpg) |
| Hesseltal Formation | 94–93 Ma | Saxony &North Rhine-Westphalia, Germany | Deposited during the anoxic conditions of theCenomanian-Turonian boundary event, this formation has a high number of well-preserved, articulated fish skeletons, in addition to exceptionally preservedammonites with soft parts.[176][177] |  |
| Vallecillo (Agua Nueva Formation) | 94–92 Ma | Nuevo León, Mexico | The site is noted for its qualities as akonservat-lagerstätte, with notable finds including the plesiosaurMauriciosaurus and the possible sharkAquilolamna. |  |
| Conulus Bed | Turonian | Poland | A crinoid konzentrat-lagerstätte.[178] | |
| Akrabou Formation (Gara Sbaa/Agoult & Goulmima) | ? (Turonian) | Asfla, Morocco | Marine site known for exceptionally preserved, three-dimensional fish fossils.[179][180] |  |
| Orapa diamond mine | Turonian | Botswana | An insect lagerstätte known for being one of the few entomofaunas from southern Africa, containing a variety of insects,[181] particularly beetles.[182][183] | |
| New Jersey amber | 91-89 Ma | New Jersey, US | Turonian-aged amber from theRaritan &Magothy Formations of New Jersey, with a high diversity of well-preserved insects, plants and fungi.[184] |  |
| Lower Idzików beds | 87-86 Ma (Coniacian) | Lower Silesian Voivodeship, Poland | An exposure of these beds nearStary Waliszów contains akonzentrat-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-preservedphosphatized decapod remains are known.[185] | |
| Smoky Hill Chalk | 87–82 Ma | Kansas andNebraska, US | ACretaceouskonservat-lagerstätte known primarily for its exceptionally well-preservedmarine reptiles. Also known from this site are fossils of largebony fish such asXiphactinus,mosasaurs, flying reptiles or pterosaurs (namelyPteranodon), flightless marine birds such asHesperornis, andturtles. |  |
| Ingersoll Shale | 85 Ma | Alabama, US | ALate Cretaceous (Santonian) informalgeological unit in eastern Alabama. Fourteen theropod feathers assigned to birds and possiblydromaeosaurids 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, olderSannine Lagerstätte, but in a deep-water environment. Includes a high number of well-preserved shark body fossils, in addition to cephalopods and deepwater arthropods.[173] |  |
| Calcare di Aurisina | ~80-70.6 Ma | Italy & Slovenia | A late cretaceous shallow marine series of carbonate platforms dominated byrudists, with fossils of invertebrates and vertebrates, specially fishes.[186] Its best known outcrop is the Villaggio del Pescatore site, that yielded the holotype ofTethyshadros, as well other exceptionally preserved taxa likeAcynodon adriaticus.[187][188] |  |
| Auca Mahuevo | 80 Ma | Patagonia, Argentina | A Cretaceouslagerstätte in the erodedbadlands of thePatagonian province ofNeuquén, Argentina. The sedimentary layers of theAnacleto Formation at Auca Mahuevo were deposited between 83.5 and 79.5 million years before the present and offers a view of a fossilizedtitanosaurid nesting site. |  |
| Ellisdale Fossil Site | 79-76 Ma | New Jersey, US | A middleCampaniankonzentrat-lagerstätte from theMarshalltown 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 aremicrofossils.[189] | |
| Coon Creek Formation | 76.8-76.0 Ma[190] | Tennessee andMississippi, 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.[190][191] |  |
| Baumberge Formation | ~75-72 Ma | North Rhine-Westphalia, Germany | A Late Campanian formation in theBaumberge of Germany with a high number of articulated fossil fish remains, in addition to shark body fossils.[176][192] |  |
| Nardò (Calcari di Melissano)[193] | ~72-70 Ma (upper Campanian-lowerMaastrichtian) | Apulia, Italy | This site is especially famous for its limestones containing abundant fossil fish remains. |  |
| Harrana (Muwaqqar Chalk Marl Formation) | 66.5-66.1 Ma (LateMaastrichtian)[194] | Jordan | Phosphatic deposits formed in this site are known to preserve vertebrate fossils with soft tissue, such asmosasaurs, plesiosaurs, sharks, bony fish, turtles and crocodylians.[195] |  |
| Tanis[196] | 66.0 Ma | North Dakota, US | Tanis is part of the heavily studiedHell Creek Formation, a group of rocks spanning four states in North America renowned for many significant fossil discoveries from the Upper Cretaceous and lowerPaleocene. 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 giantChicxulub asteroid in extreme detail. This impact, which struck theGulf 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). |  |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Baunekule Facies | 64-63 Ma (Danian) | easternDenmark | These facies of theFaxe Formation document an extremely well-preservedcold-water coral mound ecosystem dominated byDendrophyllia corals, and also includes gastropods, tubeworms, bivalves, bryozoans and gastropods.[197] | |
| Tenejapa-Lacandón Formation | ~63 Ma | Chiapas, Mexico | A formation with a high number of well-preserved fish fossils indicative of mass mortality events.[198] 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.[199] | _Figure_4.jpg) |
60 Ma | Auvergne, France | A Palaeocene maar lake containing three-dimensional plant remains.[200] It is particularly notable for preserving one of the oldest known bees.[201] |  | |
| Danata Formation | 56-53 Ma | westernTurkmenistan | Outcropping in theKopet Dag range, this formation preserves numerous fossil fish from a northeastern arm of theTethys Ocean during thePaleocene-Eocene thermal maximum. 38 taxa from 13 orders are known, the vast majority of which areacanthomorphs.[202][203] |  |
55–53 Ma | Preserves abundantfossil fish, insects, reptiles, birds and plants. The Fur Formation was deposited about 55 Ma, just after thePalaeocene-Eocene boundary, and its tropical or sub-tropical flora indicate that the climate after thePaleocene-Eocene Thermal Maximum was moderately warm (approximately 4-8 degrees warmer than today). |  | ||
54–48 Ma | England, UK | Collected for close to 300 years, Plant fossils, especially seeds and fruits, are found in abundance. | .jpg) | |
52 - 48 Ma | British Columbia, Canada &Washington, USA | IncludesMcAbee Fossil Beds,Princeton chert &Klondike Mountain Formation; Recognized as temperate/subtropical uplands right after thePaleocene–Eocene Thermal Maximum and spanning the Early Eocene Climatic Optimum, preserves highly detailed uplands lacustrine fauna and flora. |  | |
| 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 abathypelagic habitat, forming one of the few known lagerstätte to preserve a deep-sea ecosystem.[204] | |
50 Ma | Colorado/Utah/Wyoming, US | An Eocene aged site that is noted for the fish fauna preserved. Other fossils include thecrocodilians,birds, andmammals. |  | |
50-49 Ma | Verona, Italy | A fossil site with specimens of fish and other organisms that are so highly preserved that theirorgans 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.[205] |  | |
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 aUNESCO 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. |  | |
| Baltic amber | 47-35 Ma (Lutetian toPriabonian) | Pomeranian Voivodeship, Poland &Kaliningrad Oblast, Russia | The largest amber deposit on Earth, this amber is part of thePrussian 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.[206][207][208][209] | .jpg) |
| Kishenehn Formation | 46.2 Ma | Montana | A Middle Eocene site preserving exquisitely detailed insect specimens in oil shale.[210] | |
| Mahenge | 46 Ma | Tanzania | A terrestrial Middle Eocene lagerstätte preserving plant and arthropod fossils.[211][212] | |
45-25 Ma | Occitania, France | This site qualifies as alagerstätte because beside a large variety of mammals, birds, turtles, crocodiles, flora and insects, it also preserves the soft tissues of amphibians andsquamates, in addition to their articulated skeleton in what has been callednatural mummies. |  | |
| Na Duong | Priabonian | Northern Vietnam | A Late Eocene site notable for its highly detailed coprolite preservation.[214] | |
| Bitterfeld amber | 38-34 Ma | Saxony, Germany | One of three major Paleogene deposits of European amber, this deposit of theCottbus Formation shares a similar biota to the Baltic Amber, indicating a concurrent formation, but appears to have a geologically distinct origin from Baltic amber.[209] | _Figure_4.jpg) |
| Rovno amber | 38-34 Ma | Rivne Oblast,Ukraine &Gomel,Belarus | One of three major Paleogene deposits of European amber, this deposit of theObukhov 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.[206] |  |
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 ofSequoia affinis |  | |
| Babaheidar (Pabdeh Formation) | late Eocene/early Oligocene | Chaharmahal and Bakhtiari province, Iran | A well-preserved marine fossil site in the Zagros Mountains with thousands of known fish fossils, as well as plants, crustaceans, insects, and birds.[215] |  |
| Canyon Ferry Reservoir | 32.0 ± 0.1 Ma | Montana, US | A diverse Early Oligocene plant and insect fossil site.[216] | |
| Luberon (Campagne-Calavon Formation) | ~30 Ma | Cereste, France | A group of early Oligocene localities deposited along a large freshwater lake, preserving the fossils of plants, insects, fish, and terrestrial vertebrates, often with articulated skeletons, skin outlines, feathers, and original pigmentation patterns.[217] |  |
| Rauenberg/Frauenweiler Clay Pit | 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.[218] | |
| 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,[219] including mummified fruits.[220][221][222] | |
Enspel Lagerstätte | 24.79-24.56 Ma | Rhineland-Palatinate, Germany | ||
| Aix-en-Provence Formation | ~24 Ma | Provence, France | A terminal Oligocene brackish palaeoenvironment.[224] |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| Dominican amber | 30–10 Ma | Dominican Republic | Dominican amber differentiates itself fromBaltic amber by being nearly always[citation needed] transparent, and it has a higher number offossil inclusions. This has enabled the detailed reconstruction of the ecosystem of a long-vanishedtropical forest.[225] |  |
| Riversleigh | 25–15 Ma | Queensland, Australia | This locality is recognised for the series of well preserved fossils deposited from theLate Oligocene to theMiocene. The fossiliferous limestone system is located near theGregory River in the north-west ofQueensland, 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. |  |
| Foulden Maar | 23 Ma | Otago, New Zealand | These layers ofdiatomite 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,[226] along with in situ pollen.[227] | .jpg) |
| Ebelsberg Formation | 23-22 Ma | Upper Austria, Austria | ThisAquitanian-agedkonservat-lagerstätte records an exceptional fossil assemblage of an enormous number of plants, fish, marine mammals, and marine invertebrates from a section of the centralParatethys Sea.[228] |  |
| Chiapas amber | 23-15 Ma | Chiapas, Mexico[229] | 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. |  |
| 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 rapidlyoxidizing in air. It has been reported that scientists have managed to isolate small amounts ofancient DNA from fossil leaves from this site. However, other scientists are skeptical of the validity of this reported occurrence of Miocene DNA. |  |
| Barstow Formation | 19–13.4 Ma | California, US | The sediments arefluvial and lacustrine in origin except for nine layers ofrhyolitictuff. It is well known for its abundant vertebrate fossils including bones, teeth andfootprints. The formation is also renowned for the fossiliferousconcretions in its upper member, which contain three-dimensionally preserved arthropods. |  |
| 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. |  |
| Morozaki Group[230] | 18-17 Ma | Aichi Prefecture,Japan | Known from well-preserved deep sea fauna including fish,starfish and arthropods likecrabs,shrimps and giantamphiopods. | |
| Sandelzhausen | 16 Ma | Bavaria, Germany | A Middle Miocene vertebrate locality.[231] | |
| 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.[232] | |
| Dolnja Stara | ~15 Ma | Slovenia | A barnacle fossil site preserving barnacles shortly after settlement attached to mangrove leaves.[233] | |
| Pisco Formation | 15-2 Ma | Arequipa &Ica, Peru | Several specialists consider the Pisco Formation one of the most importantlagerstätten, based on the large amount of exceptionally preserved marine fossils, including sharks (most notablymegalodon),penguins,whales,dolphins, birds, marine crocodiles and aquaticgiant sloths. |  |
| Hindon Maar | 14.6 Ma | New Zealand | Amaar preserving a Southern Hemisphere lake-forest ecosystem, including body fossils of plants, insects, fish, and birds,[234] along with in situ pollen[227] and coprolites of both fish and birds.[234] | |
| 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.[235] |  |
| 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.[236] | |
| 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. Newmarsupial lion,thylacine, anddasyurid material has also been recovered. | |
| Tunjice | ? (Middle Miocene) | Slovenia | This site is known worldwide for the earliest fossil records ofseahorses.[237] |  |
| Ashfall Fossil Beds | 11.83 Ma | Nebraska, US | The Ashfall Fossil Beds ofAntelope County in northeasternNebraska are rarefossil sites of the type calledlagerstätten that, due to extraordinary local conditions, capture an ecological "snapshot" in time of a range of well-preserved fossilized organisms. Ash from aYellowstone hotspot eruption 10-12 million years ago created these fossilizedbone beds. |  |
| 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. |  |
| Saint-Bauzile | 7.6-7.2 Ma | Ardèche, France | A Late Miocene site preserving articulated mammal skeletons with skin and fur impressions.[238] | |
| Capo San Marco Formation | ~7 Ma | Sardinia, Italy | A microbialite containing exceptionally preservedGirvanella-type filaments.[239] | |
| Tresjuncos | 6 Ma | Cuenca, Spain | ALate Miocene lacustrine konservat-lagerstätte containing fossils of diatoms, plants, crustaceans, insects, and amphibians.[240] | |
| Gray Fossil Site | 4.9-4.5 Ma | Tennessee, US | As the first site of its age known from theAppalachian region, the Gray Fossil Site is a unique window into the past. Research at the site has yielded many surprising discoveries, including new species ofred panda,rhinoceros,pond turtle,hickory tree, and more. The site also hosts the world's largest known assemblage of fossiltapirs. |  |
| Willershausen | Late Pliocene | Lower Saxony, Germany | A lacustrine fossil site containing well preserved beetles.[241] |
| Site(s) | Age | Location | Significance | Notable fossils/organisms |
|---|---|---|---|---|
| The Mammoth Site | 26Ka | South Dakota, US | The facility encloses a prehistoricsinkhole that formed and was slowly filled withsediments during thePleistocene era. As of 2016, the remains of 61mammoths, including 58 North AmericanColumbian and 3woolly mammoths had been recovered. Mammoth bones were found at the site in 1974, and a museum and building enclosing the site were established. |  |
| RanchoLa Brea Tar Pits | 40–12 Ka | California, US | A group oftar pits where naturalasphalt (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 arePleistocene mammoths,dire wolves,short-faced bears,American lions,ground sloths, and, thestate fossil ofCalifornia, the saber-toothed cat (Smilodon fatalis). |  |
| Waco Mammoth National Monument | 65–51Ka | Texas, US | A paleontological site andmuseum inWaco, Texas,United States where fossils of 24 Columbian mammoths (Mammuthus columbi) and other mammals from the PleistoceneEpoch have been uncovered. The site is the largest known concentration of mammoths dying from a (possibly) reoccurring event, which is believed to have been aflash flood. |  |
| 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 liketoxodonts,glyptodonts,camelids, and thefelidHomotherium venezuelensis. |  |
| 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 | 500-1Ka | South Australia, Australia | A series of caves that preserve numerous pleistocenemegafauna, likeThylacoleo, and is recognized as a World heritage site alongside the older, but geographically similar Riversleigh site. |  |
| 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 famousdodo. The mortality assemblages may have formed from a freshwater lake that was occasionally impacted by catastrophic droughts.[242] |  |
| Crawford Lake | 800Ya-present | Ontario, Canada | This lake is notable for its detailed preservation ofrotifer anddinoflagellate fossils even after centuries, documenting the ecological shifts that occurred in and around the lake following the establishment of aIroquoian village from 1268–1486 CE, and later following European colonization of the region in the early 19th century.[243] |
The identity of the [Francevillian biota] macrostructures remains unknown and their biogenicity is questionable
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