List of Biomaterial Fossil Papers (maintained)
Peer-reviewed journal articles on surviving endogenous biological material including tissue and DNA.DOI: 10.13140/
Brian Thomas, Ph.D. paleobiochemistry (U. of Liverpool)
Bob Enyart, Real Science Radiorsr.orgT.
Newly added papers & biomaterial dated 250M+ years in red.
#YearDescription (gray cells for dinosaur remains)AgeAuthors, Publication, etc.LinkNotes / Excerpts (papers do not qualify for this page if they report only soft tissue imprints or permineralization; they must document extant endogenous biological materal) Vol (Issue), pp, DOI, etc.Dinosaur papers:
1162021Parts of proteins in titanosaur eggshell~70MYDhiman et al. Protein fragments in late Cretaceous eggshell,"...we provide incontrovertible evidence for the preservation of proteinaceous moieties in Maastrichtian dinosaur eggshell using pyrolysis-GC9GCTOFMS."7/13/2021:1-11dino
1152021Nerve fragments, sheath, from Triceratops bone65MYArmitage, First Report of Triceratops Occipital Condyle Nerves, Microsc"In this study we provide evidence of nerve fragments, characterized by a double helical wrapping of collagen fibers, from a Triceratops condyle collected at the Hell Creek Formation, MT."29(2):20-25dino
1142020Dino eggshell peptide bonds and chelating ligands200MYNorell, Wiemann, et al. The first dinosaur egg was soft,"Hard eggshells preserve PFPs still-intact peptide bonds and contain abundant chelating ligands"; PFP, protein fossilization products, is a term Wiemann, Yang, & Norell introduced in 2018583:406–410dino
1132020Hypacrosaurus cartilage; apparent chromosomes75MYBailleul, Schweitzer et al. Evidence of proteins, chromosomes, DNA, NS"Isolated [hadrosaur] chondrocytes... positive response to two DNA assays"; "Collagen II is not produced by microbes"; In supplemental material: "chromosomes are very clear"
1122020Triceratops blood clots, veins, valves, nerve fibers65MYArmitage, Solliday, Dinosaur blood clots in vessel canals, Microscopy"dinosaur neurovascular bundles travel as a triad of veins, vessels, and nerve fibers, as in other vertebrates." Re: Schweitzer and heme, "free iron was unavailable, at least in these... bones"28(5):30-38 Sept 2020 dino
1112019Permian marker plant's lignin biopolymers ~255MYTewari et al. Biomarker signatures of Glossopteris plant. Palaeogeography, Part B
1102019T. rex cortical bone blood vessel structures66MYBoatman, Schweitzer et al. Tissue and protein preserved in T. rex, N Sci title: "Mechanisms of soft tissue and protein preservation in Tyrannosaurus rex." Preemptive criticism:
1092019Precambrian chitin (from glucose) from fungi walls ~950MYLoron et al. Early fungi from the Proterozoic era in Arctic Canada,
1082019Partially unracemized feather amino acids in amber99MYMcCoy, Collins et al. Amino acids from feathers in amber, Scientific"Burmese amber sample shows less racemisation than the Baltic amber sample which is unexpected given that Burmese amber ~99 Ma is approximately twice as old as Baltic amber ~44 Ma"9(6420):1-8
1072019Sugars from Cretaceous fungi in mudstones & coal ~25-70MYMarynoswki et al. Indicators of fungal metabolism, Int'l J of Coal"co-occurrence of trehalose, mannitol and arabitol in the sedimentary rocks... suggests their fungal origin, because these... are major compounds present in most modern fungi"May 2019, pp 51-61
1062019Lebanon's hagfish fossil with skin protein remnants ~100MYMiyashita, et al. Hagfish from the Cretaceous Tethys Sea, Proc. Nat'l Acad"The chemical mapping is consistent with the fact that a major component of hagfish slime is tightly coiled, mucin-coated α-keratin threads"
1052019Edmontosaur osteocytes, blood vessels, collagen66MYUllmann, Nellermoe et al. Patterns of cellular preservation Cretaceous vast Edmontosaurus bonebed, each specimen yields abundant osteocytes, blood vessels, and collagen; surprisingly, there's no correlation between burial depth nor spongy or dense bone.99:1-13dino
1042019Anchiornis (small dinosaur) feather keratin160MYPan, Schweitzer et al. Direct evidence of feather molecular evolution"Keratins have [enhanced] preservation potential over other nonbiomineralized tissues." This is presumed apparently because feathers are not normally saturated in hemoglobin. See 10.1073/pnas.1815703116dino
1032018Flexible portions of skin layers, red-brown liver trace180MYLindgren, Schweitzer et al. Icthyosaur homeothermy and crypsis,"still-flexible remnants of the original scaleless skin... epidermal and dermal... cellular, sub-cellular biomolecular constituents... also possible internal organ trace... red-brown liver... in rib cage."564:359–365
1022018Bacteriochlorophyll (still pink & biological) pigments1.1GYGueneli et al 1.1-billion-year-old porphyrins, Proc. Nat'l Acad of lead researcher Assoc Prof Jochen Brocks, Australian National University, "What we’ve found is the [so far] oldest biological colour."
1012018Bivalve organics in the late Cretaceous 66MYMyers et al. Cretaceous Pinna organic matrix & shell microstructure,"interprismatic proteins are preserved with intact peptide bonds, and suggest an abundance of the amino acid glycine" (8):711-714
1002018Dickinsonia lipid biomarkers showing cholesteroids558MYBobrovskiy et al. Steroids establish Ediacaran Dickinsonia as animal,"lipid biomarkers extracted from organically preserved Ediacaran macrofossils... Dickinsonia... produced cholesteroids..." But Google: Günter Bechly dickinsonia, for his rebuttal.361(6408):1246-1249
992018Starch (a polysaccharide) in a forest deposit280MYLiu et al. Fossil starch reveals early plant–animal mutualism,"ultrathin sections of the megaspores indicate that they may have been produced after the megaspores were fertilized" and "used to attract and reward animals... for megaspore dispersal"46(5):423-426
982017Sea turtle muscle protein, beta-keratin, pigment54MYLindgren, Schweitzer et al. Juvenile sea turtle, Nature: Scientific long turtle (as with entire list, no report that it was found coated in biological iron)7(13324):1-13
972017Bird preening (sebaceous) gland still containing oil48MYO'Reilly et al. Uropygial gland lipids in 48MY-old bird, Proc. Royal Soc.
962017Dinosaur eggshell color by protoporphyrin/biliverdin66MYWiemann et al. Dinosaur egg color: oviraptors laid blue-green eggs,"Preserved dinosaur egg color... pushes the current limits of the vertebrate molecular and associated soft tissue fossil record..."PeerJ 5:e3706dino
952017Ginkgo leaves "full of organic molecules"200MYVajda et al. Molecular signatures of leaves unexpected, Nature Eco &
942017Melanin in nodosaurus dino (and other biologicals)112MYBrown et al. Exceptionally preserved armored dinosaur, Current Biology
932017Tick-related mammalian "red blood cells"20-45MYPoinar, Tick erythrocytes reveal piroplasms, Journal of Medical
922017Nothosaurus coracoid vessel, hydroxyproline~225MYSurmik et al. Intraosseous fossilized middle Triassic soft tissue, Sci of
912017Reconfirmation of B. canadensis dinosaur collagen80MYSchroeter, Schweitzer et al. Expansion of sequence, J of Proteome
902017Lufengosaurus rib collagen195MYLee et al. Collagen in an early Jurassic sauropod, Nature
892016Proteinaceous amide groups in chert1.88GYAlleon, Bernard et al. Molecular preservation of 1.88 Ga... organic...,
882016Early Triassic reptile blood vessels, etc.247MYSurmik et al. Organic Matter from Triassic Reptile, PLoS
872016Psittacosaurus dinosaur skin now also has keratin130MYVinther et al. Camouflage in an Ornithischian dinosaur, Current–2462dino
862016Beta-keratin in oviraptor dinosaur claw75MYMoyer, Schweitzer et al. The claw of nesting dinosaur, Proc Royal Society
852015Hadrosaur "blood vessels from Brachylophosaurus"80MYCleland, Schweitzer et al. Blood vessels characterization, J Proteome first paper to sometimes drop qualifier "like" as in "blood-vessel-like" and state clearly "dinosaur vessels", etc.14(12):5252-5262dino
842015Dinosaur collagen and red blood cells75MYBertazzo, et al. Fibres and cellular structures..., Nature
832015Dinosaur melanosomes and pigment150MYLindgren et al. Molecular composition of Jurassic feathers, Scientific
822015A pterosaur's orange claw material150MYHone et al. A specimen of Rhamphorhynchus with soft tissue,
"The claw sheathes seen here do seem to be genuinely preserved soft tissues given their clear colour and texture differences to the surrounding matrix."
PeerJ 3:e1191
812014Precambrian multicellular animals (metazoans)530MYMoczydłowska et al. Biogeochemisty of Ediacaran Metozoa, J of
802014Bird, marine reptile, mosasaur, icthyosaur pigments55-190MYEdwards et al. Pigments through time [survey], Pigment Cell Melanoma" far the data show that original chemical retention may be more common than previously believed."27(5):684-685
792014Turtle, mosasaur, and icthyosaur eumelanin55-190MYLindren et al. Pigmentation... melanism in extinct marine reptiles,
782014Gastropod egg capsule chitin200MYWysokowski et al. Chitin in 200-million-year-old egg capsules,'s an egg capsule? See Wikipedia for "Otheca" and shark cases
772014Extinct aquatic bird Gansus feather melanosomes~125MYMoyer, Schweitzer et al. Melanosomes or Microbes, Scientific"have refined the age of the Xiagou Formation lacustrine beds to the early Aptian stage"srep04233
762013Triceratops osteocytes; soft sheets of fibrillar bone 65MYArmitage, Anderson, Soft sheets of horn fibrillar bone, Acta in Italy (Acta Histochemica)Hear interviews with the authors
752013Vauxia sea sponge chitin505MYEhrlich et al. 505-million-year old chitin in Vauxia gracilenta, Scientific
742013Lufengosaurus embryo bone protein190MYReisz, Embryology of early Jurassic dino with evidence of organic...
False: Earliest 'known' organic remains of a terrestrial vertebrate. See 1966 below.
732013Signature of blood in mosquito gut46MYGreenwalt et al. Hemoglobin-derived porphyrins. Proc. Nat'l Acad of
722013Dino DNA & proteins actin, tubulin, PHEX, histone66-80MYSchweitzer et al. Analyses of dino osteocytes support endogenous..."All of our assays require at least 4 or more bases to generate reactivity... each capitalizing on different aspects of the chemistry of DNA... [show] identical patterns..."52(1):414-23dino
712013Biological material in crinoids350MYO'Malley et al. Taxon-specific organic molecules,
702012Cuttlefish ink sac160MYGlass et al. Jurassic period eumelanin pigment, Proc. Nat'l Acad. of
692012Archaeopteryx black melanin organosulpher residue150MYCarney et al. Color and nature of Archaeopteryx feather,"not due to precipitation of the inorganic mineral"ncomms1642
682012Jurassic turtle osteocytes145MYCadena, Schweitzer, Osteocytes preservation from the Jurassic,
672011Chitin arthropod (scorpion) and eurypterid proteins417MYCody et al. Molecular signature of Paleozoic chitin-protein complex,
662011Type 1 collagen in Cretaceous mosasaur humerus~70MYLindgren et al. Microspectroscopic evidence of bone proteins, PLoS "4.68%±0.1 of modern 14C" attributed to contamination yet Nature 2017 on "Carbon fixation" finds cynobacteria get "virtually all" its carbon from the bone; not the atmosphere; e19445
652011Endogenous Mississaepia cuttlebone chitin sheets34MYWeaver et al. Late Eocene cuttlefish B-chitin consistent organics, PLoS
642011Lizard tail skin breakdown products50MYEdwards et al. Tissue preservation in 50MY-old reptile skin, Proc Royal Soc
632011Type l collagen in T. rex and hadrosaur dinosaurs68MYSan Antonio, et al, Dinosaur peptides... mech of protein survival, PLoS
622011Bird feather pigment from Confuciusornis sanctus120MYWogelius et al. Biomarkers for eumelanin pigment in fossil record,
612010Stegosaurus plate keratin~150MYChristiansen et al. Stegosaur impressions, Swiss Journal of
602010Messel feather highly reflective nanostructure40MYVinther et al. Structural coloration in a fossil feather, Roy Soc Biology
Published online Aug. 26, 2009. "The keratin... has degraded... the preserved layer of melanosomes is apparently highly reflective."
592010Archaeopteryx feather shaft original chem remains150MYBergmann et al. Archaeopteryx feathers & bone chemistry, Proc Nat Acad
582010Sinosauropteryx compsognathid dino melanosomes125MYZhang et al. Fossilized melanosomes and the colour of dinos & birds,
572010Hadrosaur biomolecules, collagen & other proteins80MYPeterson et al. Microbial biofilms [&] soft tissue in archosaurs, PLoS"compelling evidence countering the argument that these structures are simply microbial biofilm."pone.0013334dino
562010Psittacosaurus dinosaur skin color125MYLingham-Soliar et al. Integument & color of a ceratopsian, Science of in Germany (Naturwissenschaften) sample used in PRSB 275:775-780.97:479-486dino
542010Mammal hair in amber100MYVullo et al. Mammalian hairs in early Cretaceous amber, Science of
532009Hadrosaur vessel proteins, sequenced collagen80MYSchweitzer et al. Biomolecular... protein sequences of hadrosaur,
522009Hadrosaur skin cell structures/macromolecules66MYManning et al. Chemistry in a mummified hadrosaur, Proc. Royal Society Hadrosaur their teeth were more advanced than today's grazers.
512009Salamander muscle, whole18MYMcNamara et al. Organic... fossil musculature with ultracellular detail, P RS
502008Feather melanocytes100MYVinther et al. The color of fossil feathers, Royal Society Biology
492008Psittacosaurus skin fibers~125MYLingham-Soliar, Psittacosaurus complex fibre architecture, Proc Royal Soc
482008T. rex amino acid sequences68MYSchweitzer et al. Molecular methods examining..., Comptes Rendus
472008Osteocyte (bone cells)5MYBell et al, Mineralized osteocyte: a living fossil, American Journal of
462007Tyrannosaurus rex collagen68MYSchweitzer et al. T. rex soft tissue suggests the presence of protein,
452007Triceratops blood vessels, osteocytes66MYSchweitzer, Horner et al. Cellular preservation from Cretaceous, Proc RS Table 1274(1607):183-197dino
442007T. rex proteins sequenced68MYAsara, Schweitzer et al. Mastodon & T. rex protein sequences,
432007T. rex proteinaceous materials, sequenced collagen40-150MYSmejkal, Schweitzer, Will tech. enable dino proteomics, Exp Rev
422006Frog bone marrow10MYMcNamara et al. Organic preservation of amphibian bone marrow, 2006 also, Schweitzer's Dangerous Discovery, dinosaurs smell like
412005Sauropod egg shell ovalbumin protein~75MYSchweitzer et al. Molecular preservation in dinosaur eggshells, P Roy Soc
402005Avian bone collagen implying "molecular integrity"~75MYAvci, Schweitzer et al. Bone collagen from late Cretaceous,
"No appreciable differences in collagen pull-off properties were measured between fossil and extant bone samples under physiological conditions."
392005Dinosaurian cellular and sub-cellular preservation~68MYWittmeyer, Schweitzer, Dino vessels and cells in fossils, J Vert. 65th Ann. Mtgdino
382005T. rex vessels, cells, soft flexible connective tissue68MYSchweitzer, Horner et al. T. rex soft-tissue vessels, cellular preserv.,
372004Isolated non-spore-forming bacteria living in amber 120MYGreenblatt et al. Micrococcus luteus - Survival in Amber, Microbial
362003Skin fibers130MYE. Frey et al., Palaeobiology of Pterosaurs, Geological Society 217
352003Algal and bacterial organic molecules2.5GYBrocks et al. Biological diversity in molecular fossils, Geoch Cosmochem
342003Iguanodon dinosaur osteocalcin protein120MYEmbery et al. Iguanodon bone proteinaceous material, Connective Tissue the full paper
332002Metasequoia cellulose (with no O18)34-56MYJahren, Sternberg, Oxygen isootopes in Arctic fossil wood, GSA
322001Tyrannosaur rex collagen SEM scans65MYArmitage, Mummified collagen fibers in fossil T. rex bone, CRS
312000Live bacteria from halite [salt] deposit250MYVreeland et al. Isolation of halotolerant bacterium from salt crystal, protocol
302000Non-collagen dinosaur proteinaceous extracts125MYEmbery et al. Non-collagenous proteins from dinosaur Iguanodon, Conn Tis"guanidinium chloride extractable protein [with the] levels of glycine, aspartate and serine tentatively suggest the presence of phosphoproteins."41(3):249-259dino
291999Alvarezsaurus dinosaur β-keratin protein ~100MYSchweitzer et al. Beta-keratin immunological reactivity, J Exper.
281999Similar live bacteria strains from separate rock salts250MYStan-Lotter et al. Halococcus in Permo-Triassic salt deposits,
271999Keratin in Madagascar bird65MYSchweitzer et al. Immunoreactivity in Cretaceous bird, J. of Vert.
261998Tarbosaurus (T. rex) blood vessels80MYPawlicki et al. Blood vessels and red blood cells in dinosaur, Annals in Germany (Anatomischer Anzeiger)180(1):73-77dino
251997T. rex peptides and proteins; likely type 1 collagen~68MYSchweitzer et al. Biomolecules in Cancellous Bone, J. of Vert.
241997Fossil diatom chitin24MYStankiewicz et al. Preservation of chitin in 25-million-year-old fossils,
231997Heme in Tyrannosaurus rex bone67MYSchweitzer et al. Heme compounds in dinosaur trabecular bone,
221995Live bacteria spores from a bee encased in amber25-40MYCano and Borucki, Revival of bacterial spores in Dominican amber, didn't clone but "revived" this bacteria, in