라거슈테테

3. 화석의 특수한 보존 (보존 라거슈테테에서)

보존 라거슈테테는 일반적인 화석 산지와 달리 생물의 유해가 원래 모습과 가깝게 산출된다. 이는 화석화를 방해하는 요인(부패나 유해의 이동 등)이 억제된 특수한 환경에서 퇴적되었기 때문이다. 일반적인 환경에서는 생물이 죽으면 미생물이나 청소 동물에 의해 부패되거나, 물살이나 폭풍 등에 의해 흩어지고 부서지지만, 보존 라거슈테테는 이러한 요인들이 억제된 환경에서 만들어진다.

3. 1. 퇴적 양상

3. 2. 보존 양상

• 버지스셰일형 보존(Burgess Shale-type preservation): 캄브리아기에 아주 흔하고, 그 이후에는 드문 보존 양상이다. 생물의 유해가 분해되지 않고, 얇은 케로겐의 막이 되어 보존된다. 퇴적된 후, 물과 닿는 면에 생긴 탄산염 자생광물이 산소와 미생물의 유입을 막아서 부패가 되지 않았다고 생각된다.
• 오슈텐형(Orsten-type preservation): 미세한 절지동물의 큐티클층이 인산염이나 규산염으로 치환되어, 원래의 입체적인 모습이 잘 남아있다. 석회암에 퇴적된 것들은 산을 이용해 석회를 녹여 그 안의 미세한 화석만을 따로 떼어낸다.
• 에디아카라형 보존(Ediacara-type preservation): 에디아카라기 시대에 흔하던 끈적한 미생물 막에 생물의 유해가 도장처럼 찍혀 그 모습이 화석으로 남는다.

• 인산화: 연조직을 인산염 광물로 대체 (예: 오르스텐형 및 두샨퉈형 보존)
• 규화 작용: 연조직을 실리카로 대체하거나 가두는 현상 (예: 규화목 또는 비터 스프링스형 보존)
• 케로겐화: 연조직을 불활성 탄소질 막으로 변환 (예: 버제스 셰일형 보존에서 발견됨)
• 알루미노규산염화: 연조직을 알루미노규산염 광물 막으로 대체하거나 코팅하는 현상
• 황철석화: 연조직을 황철석으로 대체 (예: 비처 삼엽충형 보존에서 발견되는 정교한 디테일)
• 석회화: 연조직을 방해석 광물로 대체하는 현상
• 사이드라이트 또는 방해석 결절: 부패하는 연조직에서 방출된 화학 물질이 주변 퇴적물을 사이드라이트 응결 또는 화석을 둘러싸는 석탄 공으로 변형 (예: 메이슨 크릭 화석층에서 발견됨)
• 급속한 퇴적물 시멘테이션: 주변 퇴적물에 주형 및 몰드를 통해 보존된 연조직의 흔적 (예: 미생물 매트에 의해 촉진된 에디아카라형 보존)
• 호박: 경화된 나무 수지에 갇힌 연조직

4. 라거슈테테의 사례

• 인산화 (연조직을 인산염 광물로 대체, 예: 오르스텐형 및 두샨퉈형 보존)
• 규화 작용 (연조직을 실리카로 대체하거나 가두는 현상, 예: 규화목 또는 비터 스프링스형 보존)
• 케로겐화 (연조직을 불활성 탄소질 막으로 변환, 예: 버제스 셰일형 보존에서 발견됨)
• 알루미노규산염화 (연조직을 알루미노규산염 광물 막으로 대체하거나 코팅하는 현상)
• 황철석화 (연조직을 황철석으로 대체, 예: 비처 삼엽충형 보존에서 발견되는 정교한 디테일)
• 석회화 (연조직을 방해석 광물로 대체하는 현상)
• 사이드라이트 또는 방해석 결절 (부패하는 연조직에서 방출된 화학 물질이 주변 퇴적물을 사이드라이트 응결 또는 화석을 둘러싸는 석탄 공으로 변형, 예: 메이슨 크릭 화석층에서 발견됨)
• 급속한 퇴적물 시멘테이션 (주변 퇴적물에 주형 및 몰드를 통해 보존된 연조직의 흔적, 예: 미생물 매트에 의해 촉진된 에디아카라형 보존)
• 호박 (경화된 나무 수지에 갇힌 연조직)

장소	지질 시대	위치	중요성	주목할 만한 화석/생물
라이니 처트	400Ma 전	스코틀랜드, 영국	식물, 곰팡이, 지의류, 동물(유티카르키노이드, 새우, 거미류, 곤충 등)이 화산 퇴적물과 온천에 의해 제자리에 보존. 최초로 알려진 완전 육상 생태계 중 하나.
왁스바일러 라거슈테테 (Klerf Formation)	409Ma~392Ma 전	아이펠, 독일	잘 보존된 협각류, Jaekelopterus rhenaniae의 거대한 발톱 화석으로 유명. 지금까지 알려진 가장 큰 절지동물.
Heckelmann Mill	395Ma 전	라인란트팔츠, 독일	늦은 엠스절의 매우 풍부한 두족류 화석과 함께 잘 보존된 리노카리디드 고생물연갑류 보존.[50]
훈스뤼크 슬레이트 (Bundenbach)	390Ma 전	라인란트-팔츠, 독일	연조직이 보존된 몇 안 되는 해양 데본기 라거슈테테 중 하나. 많은 경우 화석은 황철석 표면층으로 덮여 있음.
고고 형성	380Ma 전 (프라니안)	서호주	신경 및 탯줄과 같이 섬세한 조직의 3차원 연조직 보존. 50종 이상의 어류와 절지동물 기술.
미구아샤 국립공원 (Escuminac Formation)	370Ma 전	퀘벡, 캐나다	희귀하고 고대 종인 물고기, 동물군, 포자 화석 발견. Eusthenopteron은 초기 사지형류와 특징을 공유하는 육기어류.
코왈라 라거슈테테	~368Ma 전	Świętokrzyskie Voivodeship, 폴란드	무기질화되지 않은 조류와 절지동물의 화석으로 알려진 후기 데본기 지층.[52]
마이더 분지	368Ma 전 (Thylacocephalan Layer)	안티 아틀라스, 모로코	연골어류와 판피어류를 포함하여 파메니안 동물군의 잘 보존된 화석 제공. 연조직도 보존.[53]
스트루드[54]	? (후기 파메니안)	나뮈르주, 벨기에	주로 어린 판피어류가 알려져 있으며, 판피어류의 보육장소임을 시사.[55] 사지형류, 절지동물, 식물 등 다양한 생물군. Strudiella는 가장 초기의 곤충일 수 있지만, 친연성은 논쟁의 여지가 있음.
Canowindra, 뉴사우스웨일스 (Mandagery Sandstone)	360Ma 전	호주	우연히 발견. 육기어류와 판피어류 물고기가 예외적으로 잘 보존.
워털루 팜 라거슈테테 (Witpoort Formation)	360Ma 전	남아프리카 공화국	최신 데본기 동안 고위도(극지방 근처) 연안 생태계에 대한 유일한 기록 제공. 데본기 후기의 고위도 조건에 대한 수많은 가정을 뒤집음.

4. 1. 선캄브리아 시대

4. 2. 고생대

• 버지스 셰일 (캄브리아기, 캐나다): 버지스셰일형 보존의 가장 대표적인 예시이다. 셰일층 중에서도 월컷 채석장(Walcott Quarry)와 마블 캐니언 채석장(Marble Canyon Quarry)가 수려한 화석으로 유명하다. 캄브리아기 폭발의 생명체가 산출되며, 아노말로카리스, 삼엽충, 할루키게니아 등이 대표적이다.
• 마오톈산 셰일 (캄브리아기, 중국): 버지스셰일형 보존의 또 다른 사례로, 버지스 셰일보다 오래된 지층이다. 버지스 못지않은 수려한 화석이 발견되며, 초기 척삭동물의 화석이 유명하다.
• 시리우스 파셋 (캄브리아기, 그린란드)
• 페조우아타층 (오르도비스기, 모로코): 오르도비스기의 버지스셰일형 보존 라거슈테테이다. Storm wave base 위에서 퇴적된 산지와, 아래에서 퇴적된 산지가 있다. 캄브리아기의 끝에 멸종했다고 생각되었던 동물들이, 오르도비스기 이후로 번성한 생물들(필석 등)과 같이 발견된다^[209].
• 비처 삼엽충층 (오르도비스기, 미국)
• 라이니 처트 및 윈디필드 처트 (데본기, 영국): 700m 정도 떨어진 두 개의 동일 시대 화석 산지이다. 화산지대에서 규산질이 함유된 뜨거운 물이 분출하여 생물들이 순식간에 규화되어 보존되었다. 원시적인 관다발식물이 많이 발견되는데, 리니아(''Rhynia'')라는 종류가 대표적이다. 그 사이에 살던 톡토기, 선충, 협각류 등 초기 육상동물도 발견된다.
• 메이즌 크릭 (석탄기, 미국)

4. 3. 중생대

• 포시도니아 셰일(Posidonia Shale) (쥐라기, 독일): 해저의 저산소 환경에서 쌓인 흑색 셰일이다. 거의 완벽하게 보존된 벨렘나이트와 어룡 화석, 그리고 18m짜리 통나무에 다닥다닥 붙은 바다나리 세이로크리누스(''Seirocrinus'') 군집 화석이 유명하다.
• 졸른호펜 석회암층(Solnhofen Plattenkalk) (쥐라기, 독일)
• 진주층 (백악기, 대한민국): 호수에서 퇴적된 지층으로, 무척추동물의 체화석 및 척추동물의 생흔화석이 대표적이다.
• 크라투층(Crato Formation) (백악기, 브라질)

4. 4. 신생대

5. 한반도의 라거슈테테

진주 지층은 1억 1240만–1억 650만 년 전의 지층으로, 제홀 그룹 이후의 곤충 집합체^[128]와 등각류, 어류와 같은 다른 동물군으로 유명하다.^[129][130] 이 지층은 또한 풍부하고 다양한 사지동물의 발자국으로 유명하다.^[131]

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_[52] 논문 Kowala Lagerstätte: Late Devonian arthropods and non-biomineralized algae from Poland https://onlinelibrar[...] 2023-06-29
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_[70] 논문 A volcanic tuff near the Carboniferous–Permian boundary, Taiyuan Formation, North China: Radioisotopic dating and global correlation https://linkinghub.e[...] 2021-11-01
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_[81] 논문 Palynological constraints on the provenance and stratigraphic range of a Lopingian (Late Permian) inter-extinction floral lagerstätte from the Xuanwei Formation, Guizhou Province, China https://www.scienced[...] 2023-06-18
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_[85] 논문 A probable oligochaete from an Early Triassic Lagerstätte of the southern Cis-Urals and its evolutionary implications https://www.research[...] 2023-04-28
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_[90] 논문 The Oldest Aphid Insect from the Middle Triassic of the Vosges, France https://bioone.org/j[...] 2024-10-31
_[91] 논문 Revisiting horseshoe crab fossils from the Middle Triassic (Anisian) Strelovec Formation Konservat-Lagerstätte of Slovenia https://rune.une.edu[...] 2021
_[92] 논문 Paleoecology, Taphonomy, and Biogeography of a Coenothyris Community (Brachiopoda, Terebratulida) from the Triassic (Upper Anisian–Lower Ladinian) of Israel http://www.bioone.or[...] 2005
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_[94] 논문 Bromalites from the Upper Triassic Polzberg section (Austria); insights into trophic interactions and food chains of the Polzberg palaeobiota 2020-11-25
_[95] 논문 Mineralized belemnoid cephalic cartilage from the late Triassic Polzberg ''Konservat-Lagerstätte'' (Austria)
_[96] 논문 The Upper Triassic Polzberg palaeobiota from a marine Konservat-Lagerstätte deposited during the Carnian Pluvial Episode in Austria 2021-08-17
_[97] 논문 Diagenetic Alteration of Triassic Coral from the Aragonite Konservat-Lagerstatte in Alakir Cay, Turkey: Implications for Geochemical Measurements https://pubs.geoscie[...] 2013-07-15
_[98] 논문 ''Squaloraja'' Riley 1833 (Holocephala: Squalorajidae) from the Lower Jurassic of Osteno Konservat-Lagerstätte (Como, NW Italy) 2023-05-19
_[99] 논문 New records of the late Pliensbachian to early Toarcian (Early Jurassic) gladius-bearing coleoid cephalopods from the Ya Ha Tinda Lagerstätte, Canada https://onlinelibrar[...] 2023-04-08
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_[102] 논문 Taphonomy of the Lower Jurassic Konservat-Lagerstätte at Ya Ha Tina (Alberta, Canada) and Its Significance for Exceptional 2019-11-04
_[106] 논문 Dietary niche partitioning in Early Jurassic ichthyosaurs from Strawberry Bank 2022-09-29
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_[108] 논문 The Posidonia Shale (Lower Toarcian) of SW-Germany: an oxygen-depleted ecosystem controlled by sea level and palaeoclimate https://www.scienced[...] 2023-07-01
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_[110] 논문 Middle Jurassic Monte Fallano Plattenkalk (Campania, southern Italy): first report on terrestrial plants, decapod crustaceans and fishes https://www.schweize[...] 2014-04-01
_[111] 논문 EXCEPTIONAL ACCUMULATIONS OF STATOLITHS IN ASSOCIATION WITH THE CHRISTIAN MALFORD LAGERSTÄTTE (CALLOVIAN, JURASSIC) IN WILTSHIRE, UNITED KINGDOM https://pubs.geoscie[...] 2024-05-12
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_[114] 논문 Age and depositional conditions of the marine vertebrate concentration Lagerstätte at Gomez Farías, southern Coahuila, Mexico https://linkinghub.e[...] 2024-06-08
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_[116] 논문 Palaeoenvironment and taphonomy of a Late Jurassic (Late Tithonian) Lagerstätte from central Poland https://www.idunn.no[...] 2013
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_[119] 논문 The Early Cretaceous coastal lake Konservat-Lagerstätte of La Pedrera de Meià (Southern Pyrenees) https://revistes.ub.[...] 2023-04-17
_[120] 논문 Jumping bristletails (Insecta, Archaeognatha) from the Lower Cretaceous amber of Lebanon https://onlinelibrar[...] 2019
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_[123] 논문 Early Cretaceous polar biotas of Victoria, southeastern Australia—an overview of research to date https://www.tandfonl[...] 2018-04-03
_[124] 논문 Evidence of plant–insect interaction in the Early Cretaceous Flora from the Crato Formation, Araripe Basin, Northeast Brazil https://www.tandfonl[...] 2024-05-21
_[125] 논문 Taphonomy of fish, invertebrates and plant remains in the first Tethyan-South Atlantic marine ingression along Cretaceous rift systems in NE-Brazil http://dx.doi.org/10[...] 2023
_[126] 웹사이트 Vertebrates from the uppermost stratigraphic sequence of the Pietraroja plattenkalk (Early Cretaceous, Southern Italy) https://iris.unito.i[...] 2024-07-12
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_[128] 논문 Cretaceous beetles of the Jinju Formation (Coleoptera): An overview of the Jinju Formation, its coleopteran diversity, and past and future research https://linkinghub.e[...] 2024-06-08
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_[133] 논문 Ammonites from the La Negra Facies (El Doctor Formation, late Albian) of the Muhi Quarry, Hidalgo, central Mexico https://www.scienced[...] 2021-11-01
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_[192] 논문 Middle–late Miocene palaeoenvironments, palynological data and a fossil fish Lagerstätte from the Central Kenya Rift (East Africa) https://pubs.geoscie[...] 2023-06-29
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_[196] 논문 The Girvanella-like remains from Messinian marine deposits (Sardinia, Italy): Lagerstätten paradigm for microbial biota? https://linkinghub.e[...] 2024-05-12
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