Paleobiology, 14(4), 1988, pp.

331-344

cause and classification Konservat-Lagerstatten:

Peter A. Allison
Abstract.-A review of the processes required for exceptional preservationof soft-bodiedfossils inhibit decay and emphasizes the importanceof demonstratesthat anoxia does not significantly early diagenetic mineralization.Early diagenesis is the principal factoramongst the complex processes leading to soft-part preservation.The development of a particularpreservationalmineral is and salinity.A new causative classification controlledby rate of burial, amount of organic detritus, of soft-bodiedfossil biotas is presented based upon fossil mineralogyand mineral paragenesis. BS6 5DS, UnitedKingdom. of QueensRoad, Bristol University Bristol, PeterA. Allison. Dept. ofGeology, Harbor 620 Harbor, address: Road,Friday Friday Laboratories, University Current of University Washington, Washington 98250 Accepted: August 10, 1988

Introduction Exceptionallypreserved soft-bodiedfossil by biotas, termed Konservat-Lagerstatten Seilacher (1970), preserve invaluable evidence of Phanerozoic metazoan diversity.For example, accordingto Sepkoski (1981: p. 39) approximately20% of marine metazoan clades are known fromonly three Paleozoic LagerBurgess Shale, Hunsriick Slate, and stdtten: Mazon Creek. of Seilacher's (1970) classification such deposits recognized two types of occurrence: deposit,and the"Konzentrat," concentration or or the "Konservat," conservation deposit. In organic remainsare concentrated the former, by sedimentologicaland biological processes which generally exclude the preservationof organic softparts. In the latter,fossilization is broughtabout by unusual depositional conditions which lead to the preservationof exceptional details such as softparts.Seilacher (1970) considered the most criticalsedimentological controls favoringexceptional preservationto be anoxia, rapid burial, early diagenetic concretiongrowth,or the occurrence of a decay-inhibitorymedium such as tar, or Seilachpermafrost, amber.More recently, er et al. (1985) refined the classificationof conservationdeposits. Obrution (rapid burial), stagnation,and cyanobacterialcoverings were considered the principal causes of exceptional preservation.These classifications highlighted the importance of depositional
c 1988 The Paleontological Society. All rightsreserved.

conditions to the development of particular levels of preservation.Thus, obrution commonlyleads to the preservationof articulated hard parts,but is rarelythe formativeagent in the preservation of soft parts. Similarly, stagnationalone will not usually preservethe of softparts;it will, however,inhibit remains and it therefore scavenging and bioturbation, promotesthe preservationof articulatedhard parts. "Preservationaltraps,"such as permafrost,tarsands, and amber, may allow the preservationof three-dimensionalsoftparts and are associated with particular levels of preservation. For instance, permafrostmay preservesofttissuein such fineconditionthat itis edible, and amberis usually characterized by three-dimensionalpreservation of small organisms which lack internal detail. Such forms of preservation, however, are extremelyrare and rarely encountered by the majorityof paleontologists. The only longmedium thatis of any termdecay-inhibitory real geological consequence is peat bogs. Examples include the preservationof Iron Age human cadavers complete with flesh (Glob 1969) and the preservation of three-dimensional plant organs (Scott 1979) and arthropods (Bartramet al. 1987) fromthe Carboniferouscoal deposits of Europe and America. The "cyanobacterial coverings" reported by Seilacher et al. in 1985 have (to date) only been reported from a handful of localities, which include the JurassicSolnhofen Limestone of Germany(Seilacher et al. 1985), the
00 0094-8373/88/ 1404-0003/$1.

332

PETER A. ALLISON

Jurassic OxfordClay ofEngland (Martill1986, relationship between depositional environ1987), and the Eocene Messel Shale of Ger- ment and life habitat. many (Wiittke1983). By farthe greatestnumActualisticexperimentation, however, has ber of Konservat-Lagerstdtten, however, are shown thatexceptionalpreservationis no inpreservedas earlydiageneticmineralgrowths. dication of duration or nature of transport The geochemical parametersfor the forma- (Allison 1986). Freshlykilled carcassesofsofttion of such minerals have received much bodied and lightly skeletized animals tumattention in recent years (see Berner 1980, bled in a rotating barrel for several hours 1981). Particular authigenic mineral species suffered nor neitherfragmentation extensive are indicative of specificdepositional condi- disarticulation. Conversely, carcasses entions such as level of pH, Eh, organiccontent, tombed in a jar of seawater were buoyed up rate of burial,salinity,and degree of oxygen- to the surfaceand disarticulated duringdecay ation. The preservationalmineralogyofa fos- to produce a carpetofskeletalfragments upon sil deposit is therefore valuable but under- the floorof the jar. Thus, it is extentof prea appreciated indication of the depositional burial decay, ratherthan nature or duration conditions leading to fossil preservation.In of transport, which governs the operation of this study I review taphonomic conditions this process. required for soft-part preservationand propose a new classificationof Konservat-Lager- Decay: rate and inhibition stdtten based upon taphonomic featuressuch The decay of organic carbon in naturalsedas degree of compaction, level of soft-part imentary sequences is best expressedin terms and preservationalmineralogy. of a half-life(Berner 1985). The duration of preservation, this half-lifeis dependent upon: a) nature of Controls on the Formation of the decomposing carbon; b) the supply of oxtten Konservat-Lagersta idants to bacteria involved in the decompoThe salient charactersthat define the oc- sition process; and c) sediment type. currenceof a particulardeformationalstrucNature of organiccarbon. -Organic carbon tureare oftenreferred as a structural to style. occurs in a complex and variable formin asIn a paleontological context,the term"pres- sociation with other elements, principally ervational style" refersto the sum total of oxygen, nitrogen,and phosphorus. Organic taphonomic charactersexhibited by a biota molecules decay at different rates according that were produced during the fossilization to their chemical formulae and molecular Those with the shortesthalfprocess. Thus, preservationalstylewould de- configuration. scribe characterssuch as degree of fragmen- lives may be considered volatiles; those that tation and disarticulation,extent of decay demonstratea degree of decay-resistance are Cellulose and chitin,for prior to mineralization,preservationalmin- termedrefractories. eralogy,and mineral form. example, belong to a group of carbohydrates known as structural and form polysaccharides Transport complex biopolymers exhibiting considerReduced transport has been cited as a con- able decay resistance. factorin the exceptional preservatributary -The depletion of poreOxidant supply. tion of the Burgess Shale (Whittington1971; water oxygen forcesbacteria in sediment to Conway Morris1979a) and theHunsriickSlate utilize a series of alternativeoxidants during (Stiirmerand Bergstrom1973). In both ex- decay (see Aller 1980; Aller and Yingst 1980; amples, the authors argued that minimal Aller and Mackin 1984; Berner1980;Westrich transportreduced the degree of disarticula- and Berner1984). The preferred consumption tion and fragmentationof soft-bodied and of these oxidants is governed by their freelightly skeletized organisms. Logically, this energyyield during respiration.In ideal cirsuggests that exceptionally preserved biotas cumstances, they are stratifiedin the sediare indicatorsofminimaltransport and there- mentprofile(Fig. 1),with those liberatingthe foreapplies certainconstraints the spatial highest free-energy to yield occurringhighest

KONSERVAT-LAGERSTATTEN

333

AEROBIC DECAY

+ (CH2O)106(NH3)16H3P04 10602 o1 06 C02 + 16 NH3 + H3P04 + 106H20

ANAEROBIC DECAY

Manganese Reduction + 212 (CH20)106(NH3)16H3PO4 MnO2 + 332 C02 + 120 H20O*438 HCO-3 + 16NH+4

+ HPO2-4 + 212M&+4

Nitrate Reduction + -.-.-7.2 C02 + 98.8HC0-3 + 16 NH+4+ 42.4N2 CH20)106(NH3)16H3PO4 84.8NO-3

+ HP024 + 49 2?

Iron Reduction + 424 Fe(OH)3+ 756C02 o862 (CH20)106(NH3)16H3PO4

HC0-3 + 16 NH+4 + HP02-4 + 424 Fe2+ + 304H20

+ (CH20)106(NH3)16H3PO4 53 S02-4

Sulfate Reduction 39 C02 + 67 HCO-3 + 16 NH+4 + HP02-4+ 53 HS+ 39H20

+ (CH20)106(NH3)16H3PO4 14 H20

Methanogenesis (Carbonate Reduction)

-39 C02 + 14 HCO-3 + 53 CH4 + 16 NH+4 + HPO2-4

bacterial reduction zones in "ideal" sediment. Reactions occurringat the top of the column Stratified yield. Reactions lower in the sediment profileontyoperate afterthe complete reduction have a higher free-energy of ions higher in the column (afterBerner 1981).
FIGURE 1.

in the sequence (Redfield 1958; Berner1981). Reactionslower in the sedimentprofile would only operate afterthe complete reductionof ions required for the more efficientones. However, the supply of these oxidants is not uniform. Nitrate-reduction methanogenand esis dominate in a freshwaterenvironment, whereas sulfate-reduction methanogeneand sis dominatein marineregimes(Malcolm and Stanley 1982). The depth to which these reactions are active in the sediment depends upon organic supply, temperature, and sediment permeability.A greater input of organic detritusincreases demand forthe ions required for anaerobic respiration.Reduced permeability impedes the transferof these ions from overlying the watercolumnto porewaters,and temperature increases the rate of decay. In all situations,the reduction zones in the chemically stratified sediment will be attenuated.In the most extremecase, the an-

oxic/oxic boundary is projected above the interface. sediment/water Prolonged exposure to oxygen can lead to the completedestruction organiccarbon in of sediment by aerobic bacteria. However, according to J0rgenson(1982, 1983) and Peck and Legall (1982), anaerobic bacteria are less efficient biodegraders and may be unable to completely break down complex biopolymers. J0rgenson(1982, 1983) suggested that completebacterialutilizationof refractory organic carbon in sediment may thereforerequire the presence of a bacterial community composed of a complete range of reducing bacteria (e.g., aerobic bacteria, nitrate-reducers,manganese-reducers, etc.). In such an environment,each group of bacteria would partially degrade the more complex molecules to produce by-productsthat other bacteria(lower down in the chemicallystratified column) could use in the respirationprocess.

334

PETER A. ALLISON

In this model, preservationof complex biopolymers would be favored in euxinic settingswhere the bacterialreductionzones are severely attenuated and the microbial food chain is broken. Aerobic respiration of organic carbon is commonly considered to be a more rapid means of biodegradation than anaerobic decay. This is because aerobic decay processes yield (Aller 1980) have a higher free-energy and may be able to support a more diverse bacterial community(J0rgenson1982, 1983). Thus, anoxia is generally considered a prerequisite forpreservationof soft-bodiedand lightly skeletized organisms (see Seilacher 1970; Seilacher et al. 1985). However, the actualisticexperimentsof Allison (1988a) have shown that the anaerobic decay of soft-bodied and lightly skeletized organisms can be extremelyrapid. The soft parts of such organisms can be considered volatiles and are easily decayed. Anaerobic decay of such tissues is slower than aerobic decay, but, in a geological sense, it is not slow enough to promote soft-part preservation. Sedimenttype. -Long-term decay inhibition of organic soft parts is an unusual occurrence and restrictedto specific taphonomic circumstances, such as low-pH peat swamps. Exceptionally preserved plant remains in calcareous concretions from fossil peats have been recorded frommany localities. Preservationis favoredby the resistance of cellulose to decay and the antibacterial qualities of tannicand fulvicacids in the peat. However, in addition to plants,a wide array of invertebrates and vertebrates are preserved in these peats. The best example is provided by the so-called "bog people" from the peats of northernGermany,Scandinavia, Britain,and Ireland (Glob 1969). These Iron Age human cadavers have been mistakenfor recentmurdervictims(Glob 1969). Many preserve cellular detail ofskin,hair,and clothing fibers. However, preservationis solely due to the tannic and fulvic acids produced by the peat that impregnate the carcasses and preserve it like tanned leather. The Middle Eocene Geiseltales Brown Coal exposed near Halle, in East Germany (Voight 1935, 1957), is an older example of this type of preserva-

tion. Preserved soft parts fromthis locality include muscle fibers and epithelial cell structure. Fossil diagenesis as Although anoxia is ineffective a preservational medium for organic soft parts, the removal of oxygen leads to widespread reducing conditions and the production of reactive ionic species (Fig. 1). This promotesthe formation earlydiagenetic mineralprecipof itates (Berner 1981) which can formduring the first few weeks of burial. Earlydiagenetic mineral formation the only way to halt deis loss (Allison 1988a). cay-inducedinformation The most commonly occurringdiagenetic minerals associated with exceptional preservation are pyrite,carbonates (such as calcite and siderite),phosphates (such as francolite and vivianite),and silica. The mineralsoccur in a varietyof formscontrolledby degree of crystallinity, crystalsize, and growth form. Mineral formis expressed taphonomicallyin the preservation soft-bodied of biotasby three modes of preservationwhich reflect different degrees of information loss. These three modes of preservation are: permineralized tissues, mineral coats or pseudomorphs,and mineral casts. -In vivo precipiPermineralized tissues. soft tation of authigenic mineral phases prior to organic decay is rare. The most frequentoccurrence of this type of preservationis that tissues such as of permineralized refractory of cellulose and chitin.The preservation more volatile structuressuch as muscle fibersrequires exceedinglyrapid mineralizationand, as a result, is frequentlyrestrictedto phosphatic preservation(referto section on phosphates) since phosphatesmayform veryearly in the diagenetic sequence of a deposit and may pre-dateotherauthigenic mineral phases such as carbonates and pyrite (Allison 1988b). coats. -The preservation soft-part of Mineral outlines is the most frequentmode of occurrence ofsoft-bodied biotas. In thiscase, invivo precipitationof mineral phases is inhibited, of and the internalstructure softpartsis destroyed.However, the organismsact as a temand a soft-part plate for mineral formation,

KONSERVAT-LAGERSTATTEN

335

outline is preserved. Examples include: the CoMiddle CambrianBurgessShale ofBritish lumbia (Conway Morris 1985); the Upper Cambrian anthraconitesfromVasterg6tland, Sweden (Muller 1985; Muller and Wallosek 1985); the pyritizedfaunas of the Ordovician Beecher's TrilobiteBed (Cisne 1973) and the Devonian HunsriickSlate (Stiirmer and Bergstr6m 1973; Stiirmer1985); and the Eocene Messel Oil Shale biota from Darmstadt in Germany(Franzen 1985). The Messel Shale contains flattenedbody outlines of amphibians,flying mammals,and birds (Franzen 1985). These outlines are formedby a permineralizedlayer of bacteria which pseudomorph the original soft parts (Wiittke1983) by promotingthe precipitation of siderite. In this case, the bacteria responsible for decay have initiated the precipitation responsible forthe mineral coat. When bacteriadie, theyundergo autolysis, whereby enzymes and otherchemicals within the bacterial cell corrode the body wall of This the microbeand cause it to self-destruct. occurs within hours or days. The presence of mineralized bacteriatherefore implies fixing of the microbes prior to complete autolysis. Mineralization must occur very rapidly after death or even during the life of the bacteria. It may even be a cause of mortality. The Orsten biota includes three-dimensional ostracodesand crustaceansenclosed in concretions of calcium carbonate (Muller de1985). Preservationincludes finesoft-part tails such as limb morphology,eyes, sex organs and possible gut traces(Muller and Wallosek 1985). The animals are preserved in calcium phosphate and can therefore freed be fromthe enclosing calcareous concretionsby treatment with acid and then examined under the SEM. Phosphate occurs as both a replacement of cuticleand as a coat of euhedral crystals.Very little is known of the mechanism of thistypeofmineralization, itmay but have been bacteriallymediated. Ennever et al. (1981) showed thatplaque bacteriaare responsible for the formation of phosphatic coats on the teethof living mammalsand that several strainsofbacteriawere capable ofprecipitatingphosphate in this way. It is therefore possible thatthe phosphate coats occur-

ring on the Orsten arthropods were precipitated by bacteria and subsequently modifiedduring diagenesis. Tissuecastsand molds. -Soft-tissue castsand molds only preserve a two- or three-dimensional tissue outline and generally involve a greaterdegree of information loss than those mineralogicalformsalready listed. Such biotas represent the lowest level of soft-tissue preservationand have formed by the early diagenetic stabilization of sediment after a period of decay, but priorto lithification and compaction. Such stabilization is most commonly a product of early diagenetic mineral growth,but may also occur in the absence of in authigenic mineralformation naturalcasting sediments such as fine sands and muds. The Late PrecambrianPound Quartzite of the Flinders Range of South Australia contains a varietyof soft-bodiedcasts which include medusoids (Wade 1968). Preservation involved the deposition of fine-grained rhythmic "casting" sands upon the carcasses shortlyafterdeath. Organic decay continued until the organisms were destroyed. The sands, being fine-grained, underwentlittleor no compaction and retained the cast formof the carcasses. The Upper Carboniferous Mazon Creek biotaoccurring withintheFrancisCreekShale of Illinois has yielded a varietyof soft-bodied fossils preserved within siderite nodules at the base of the shale (Baird 1979). Softtissues are preserved solely as casts and molds and as impressions with little or no internal detail. Preservation occurred through early around the dediagenetic mineral formation caying organisms.Although decay has comthe pletelydestroyedmostorganicstructures, lithification sedimentaround the organism of casta replica of softparts.This replica formed a void in the subsequent concretion which was later infilled with late-stage diagenetic minerals, thus forming soft-partcasts and molds. Diagenetic Classification of Konservat-Lagersta tten Since diagenetic mineral growths are the most important factorin the formation exof ceptionallypreservedfossilbiotas,itis logical

336

PETER A. ALLISON

to erect a causative classification based upon mineral paragenesis. The following classification concentrates upon the principalfactors controlling sediment geochemistry, namely rate of burial, salinity,and organic content, and how different combinationsofthese variables function as a precipitation switch, thereby leading to mineral formation.This discussion concentrates on those minerals most commonly associated with the exceptional preservation of macrofossils (pyrite, phosphates, and carbonate). Silica is a valuable source of exceptionallypreservedmicroorganismsand three-dimensional plants,but is rarelyassociated with the preservationof volatile softpartsof macrofossils. Silica is not reviewed here, but recent reviews include those of Knauth (1979), Leo and Barghoorn (1976), and Ferriset al. (1988). Certain compounds may functionas precipitation"poisons" (pore-watercompounds thatinhibitthe formation certainminerals) of and exert an as-yet unquantified effecton mineral precipitation.The existence of such compounds is acknowledged, but because there is very little data on them,there is no discussion here on their role in early diagenetic mineral formation.The following discussion concentrates on the general geochemicalparameters (pH, Eh, organiccontent, rateof burial,and depositional environment) requiredforthe mineralizationofsoft-bodied tissues. Pyrite Pyritizedsoftpartshave only been recorded froma handful of localities including the Ordovician Beecher's Trilobite Bed of New York State (Cisne 1973), the Hunsriick Slate of the Bundenbach district Germany(Bergof strom 1989; Kott and Wiittke1987; Stiirmer 1985; Stiirmerand Bergstr6m1973), and the Middle CambrianBurgessShale ofBritish Columbia (Conway Morris 1986). It has long been known that pyrite may formvery early in the diagenetic historyof a sediment; indeed, both pyriteand its precursors have been recorded in Recent sediments(Love 1967; Lein 1978) and around the bodies of living (Clark and Lutz 1980) and dead (Allison 1988a) animals. The occurrence

of authigenic pyrite in shales has been reviewed by Brettand Baird (1986), Fisher and Hudson (1985), and Hudson (1982). These workers demonstratedthat the morphology and occurrence of pyritewas stronglycontrolled by burial rate and geochemical conditions (such as organic and sulfate content) near the sediment/water interface. Authigenic pyriteformsas a result of the action of anaerobic bacteria that promote reducing conditions. In such a system,detrital iron(and bacterially reduced iron) reactswith hydrogen sulfide (resulting fromthe reduction of pore-water sulfates) to form iron monosulfidessuch as greigite and mackinawite (Berner1970,1971). These mineralsreact in turn with bacteriallyliberated hydrogen sulfideto form authigenicpyrite (Berner1984, 1985). Thus, formation sedimentarypyrite of is facilitated the availabilityoforganiccarby bon (forbacterialrespiration), iron,and porewater sulfates(forsulfatereduction) (Berner and Raiswell 1984) (Fig. 2). In most marine fine-grained clastic sediments,iron minerals and pore-watersulfatesare present in abundance and the formation pyriteis only limof ited by the availability of organic carbon. Freshwaterenvironments, however, are typically low in sulfate content and pyriteformation is thereforesulfate-limited.Euxinic marine environments,however, are characterizedby concentrations hydrogensulfide of (frombacterialsulfatereduction),thus pyrite formationhere is determined by the availability of reactive iron (Berner and Raiswell 1984). The conditions required for pyritization of lightly skeletized and soft tissues in these three(freshwater, marine,and euxinic) environmentsare discussed below. Freshwater. -Since iron is commonlyavailable in terrestrial deposits and sulfate concentrations low, pyritization dependent are is upon the sulfate bank (Berner and Raiswell 1984). Depletion of this source is controlled by rateof sedimentationand organiccontent. In regardto the preservationof softparts,we can consider two geochemical extremes. In the first case, organic contentis high and rate of burial is low. In such a systemsulfate reduction is uniform,and the pyriteformedis disseminated evenly throughout the sedi-

KONSERVAT-LAGERSTATTEN

337

ment.The zone of bacterialsulfatereduction becomes attenuated, and sulfate-reducing bacteriaaround decomposing organic matter a are starvedof sulfate.In effect, carcass cannot functionas a reducing precipitationsink since it is surroundin such an environment, ed by decaying dispersed carbon in the sedorganiccontent iment.In the second extreme, is low and burial rate is increased. In this system, widespread pyrite precipitation the throughout sedimentis inhibited,the sulbacteria around the carcass are fate-reducing not limited by sulfate supply, and pyritization can occur. Burrowingand scavenging in such an environmentwould be inhibited by rapid burial. Because of the low concentradeposits, pyrittions of sulfatein freshwater ization is rare. An example of such deposits flora South of is theLower Devonian Gosslingia Wales (Kenrick and Edwards 1988). at Normalmarine interface or be(anoxic/oxic selvedge).-In a normalmalowsediment/water rine clastic setting, detrital iron and porewater sulfatesare presentin abundance, and is pyritization only limitedby carbon content (Bernerand Raiswell 1984). We can therefore of consider the effects burial and carbon content.Ifburial rateis low, then scavengersand bioturbatingorganisms may destroydelicate required softparts. Rapid burial is therefore to preventbioturbation.Sellwood (1971) has burial can so shown thatrapid (catastrophic?) of pore-watersulfates impede the diffusion fromthe overlyingwater column thatpyrite is formation slowed (i.e., becomes sulfate-limiting).Thus carboncontentwill exertthesame controlover the preservationof softparts as it does in freshwater systems(i.e., a low carbon content favors pyritizationof soft tissues). Pyritizationof soft parts is therefore favored by rapid burial and a low organic content. waters and resultin ments occur in stratified widespread sulfatereductionof organic matterand the productionof abundant hydrogen sulfide. Pyrite formsby the reaction of hydrogen sulfide with either reduced iron or detrital iron minerals and is limited by the supply of reactive iron (Bernerand Raiswell 1984). What,then,are the ideal conditionsfor
Euxinic marine.-Euxinic marine environ-

U.

Depositional parameters (stippled area) reof quired forpyritization lightlyskeletized and soft-bodrequiresrapied organisms.Earlydiageneticpyritization id (possibly catastrophic)burial, a low organic content, and the presence of sulfates.
FIGURE 2.

pyritizationof soft parts in this type of environment?Even though such basins are anand scavengingare preoxic and bioturbation vented, they may still be swept by currents (Baird and Brett1986) which could disaggregate a decomposing carcass.Thus rapid burial Overpreservation. would stillfavorsoft-part all organic concentrationsare typicallyhigh in euxinic environments,and pyriteformation may continue during burial to great depths (Hudson 1982). In localized areas may where carboncontentis low, pyritization be promoted by the migrationof hydrogen sulfide from organic-richareas (Berner and Riaswell 1984). What, then, is the role of organics in the preservationof softparts?Since pyritizationis limited by the availability of iron, it is likely that a low (locally) organic contentwill favorpreservationof softparts. For example, if carbon contentis locally low, thenbacterialiron reductionwill be centered reupon decomposing carcasses;the resultant duced iron will rapidly reactwith migrating hydrogensulfide,and pyritewill formin and around the carcass.If,on the otherhand, car-

338

PETER A. ALLISON

mineral formationis favored. These circumstances can be exacerbatedin organic-rich sequences where further bicarbonate ions are produced by anaerobic respiration.In such a the system, carbonate/ phosphate switchis set at the defaultposition (Fig. 3) and favorscarbonate precipitation. For phosphate minerals to form, the solubility product of calcium phosphate must be exceeded. Gulbrandsen (1969) suggested thatthis may be achieved at sites of oceanic upwelling, where plankton and algae concentratephosphates. Periodic blooms and subsequent decomposition of these organisms would liberate phosphates to pore-water solution (Lucas and Prevot 1984), thereby increasing concentrations. However, this process alone is not an adequate explanation, since the decomposition of organic matter liberatesfarmore bicarbonate ions than phosphate and may even lead to the calcium carbonate solubility product Phosphates being exceeded. Benmore et al. (1983) sugPhosphatized softpartsexhibitthe highest gested that phosphate liberated by the dequality of preservation,including three-di- composition of organic carbon would be admensional preservationof softpartsand the sorbed to ferric hydroxides within the retentionof cellular morphology.Fossilized sediment. Upon the onset of anoxia, and the muscle fibersare more commonlyassociated reductionof ferric iron,phosphates would be with phosphate replacement than with any liberated to pore-watersolution at the anoxother mineralizing agent. Examples include ic/oxicboundary,therebyproducing a phosconodont animals from the Carboniferous phate concentration peak. Low burial rate(inGrantonshrimpbed of Scotland (Briggset al. creased residence time at this level) could 1983; Aldridge et al. 1986), concavocarids(bi- promotephosphatization.According to Gulvalved arthropods)fromthe Upper Devonian brandsen (1969), precipitationof phosphate of Australia (Briggs and Rolfe 1983), squid would be favoredby an increased pH. Thus, (Donovan 1983; Donovan and Crane in prep; phosphatization would be promoted in the Allison 1988c) fromthe Oxford Clay of En- alkali reducing environmentformedby the gland, crustaceans fromthe Jurassicof Italy amines and ammonia liberated by the decay (Pinna 1985), and fishfromthe Cretaceous of of proteins in a carcass (Berner 1968). HowBrazil (Martill 1988). According to Zangerl ever, Coleman (1985) has shown that phosand Richardson (1963), Zangerl (1971), and phates have a broader stabilityfield with reConway Morris (1979b), compression of de- spect to pH than do carbonatesand may even caying softtissues is mainly the resultof col- be stable in mildly acid environments.With lapse due to decomposition. Preservationof a low burial rate,sulfidesmay be utilized by three-dimensionalsoft tissues in the exam- sulfide-oxidizing bacteria,with a resultantreples cited above thereforeimplies that min- duction in pH (Fig. 4). Coleman (1985) sugeralization was initiatedprior to appreciable gested that these circumstancescould favor decay. the formationof phosphate over carbonate. In most freshwaterand marine settings, However, both models (whethertheyrequire concentrations the bicarbonateion exceed an acid or an alkali pH) require a low rate of of those of the phosphate ion, and carbonate burial. In addition, Ennever et al. (1981) doc-

bon contentis high,thenbacterially liberated reduced iron will be rapidly depleted by organic carbon dispersed throughoutthe sediment,and pyritizationof organic concentrations, such as animal carcasses, will be impeded. Early diagenetic pyritewill occur as disseminated grains and framboidsfrom the reaction of reduced iron (comparatively reactive) and hydrogen sulfide. Later in the burialhistory thesequence,pyrite of will form by the reactionof hydrogensulfidewith (less reactive) detrital iron-bearingminerals (for example, see discussion of the JurassicPosidonia Shale in Hudson 1982). Pyritization of soft parts is thereforefavored by rapid burial and a low organic content and is most likelyto form in a marine environment where sulfateconcentrations are highest (Fig. 2).

KONSERVAT-LAGERSTATTEN

339

[ORGANIC RICH SEQUENCE |

normal pore waters HCO' P04 P04 adsorbed to ferrous hydroxides

PRECIPITATION SWITCH AT DEFAULT POSITION CARBONATE PHOSPHATE

OFF

BURIAL RAPID
residence time at LOW=anoxic/oxic boundary

EDUCED

HIGH

LIBERATION OF

P04

AT ANOXIC/OXIC INTERFACE

DISSEMINATED THROUGHOUT SEDIMENT

P04

within

sediment

CONCENTRATED AT ANOXIC/OXIC INTER ACE

CARBONATE PHOSPHATE

CARBONATE PHOSPHATE

OFF

OFF

3. FIGuRE Flow diagramsummarizingthe depositional controlsupon the carbonate/phosphate switch. precipitation High organic content and low rate of burial favors phosphate formationwhilst high organic content with rapid burial favorscarbonate formation.

umented the in vivoformation apatitecrysof talsin plaque bacteria,and Allison (1987) suggestedthatsimilarbacteriamaybe responsible forconcentrating phosphate in sediment. However, it is clear thatin some cases phosphate levels only achieve a marginalconcentrationwhere the solubilityproductis barely exceeded and widespread phosphate precipitationis inhibited.Allison (1988b) described such a systemfromthe Eocene London Clay of Kent, England, where phosphate formation was inhibitedand only occurredaround phosphatic particles,such as vertebrateand arthropod fragments,which functioned as precipitationnuclei. In a marine system,authigenic phosphates

normallyprecipitateas a sedimentaryfluorapatite known as francolite.In fresh water, however, concentrations of iron ions are higher,and the moststable phosphate species is usually vivianite (Berner 1981). The fieldofsoft-part preservationforphosphatic preservationis defined by a high organic content and a low rate of burial (Fig. 4). A high organic content supplies phosphates to pore-watersolution during the decay process.Phosphatizationofsoftpartswill be favoredby a low rateof burial,as thiswill increase residence timeat the anoxic/oxicinwhere phosphate concentrations terface, may be secondarily enriched by the reduction of ferric iron (Benmore et al. 1983).

340

PETER A. ALLISON

_j~~~~~~~~~~~~~~~~~~~~~~~~~~~;
a:

LL

cr.

4~~~~~t? ~ ~

Sellwood 1971).

In a freshwater environment,.

4. Depositional parameters (stippled area) required forphosphatizationof softparts.Earlydiagenetic phosphatization requires a low rate of burial and a high organic content.
FIGuRE

FiGrEn 5.arpoitonalean parameter (stipldeariea) rei-

Carbonates Preservationof softpartsis more commonly associated with carbonate mineralization thanwithany otherauthigenicmineralphase. The carbonateoccurs eitheras concretionsor as fine-grainedbedded limestone (often referredto as Plattenkalk deposits). Concretionary carbonatespreservingsoftparts occur as siderite or calcite and include the Carboniferous biotas of Mazon Creek,Illinois, and the Montceau les mines of France. The decompositionof organic carbon in an of anoxic environmentleads to the formation the bicarbonate ion that,if produced in sufficient can quantity, reactwith calciumor iron ions presentin pore-watersto exceed the solubilityproductof calcium carbonateand produce authigenic carbonates (Raiswell 1971, 1976; Berner 1981). In a marine system,calcium is normallypresent in supersaturation with levels, and the iron reactspreferentially sulfideions to formpyrite;thus calcite is the most likely mineral phase to form (but see

qiredtfor presnervto ofsftprt8iti1cronts anit Field exetoa of brsraicroniscaraterized lbyerap-d id (pssbl dcatasophc burialgof marorganicihsequnce

irnsufioins exdced the to solubiit poduc cofe tironar carbonateandspomoate sderite prciptoiation (Beerner1981). aiwll171 97) Toeerhe quantityen bitilcarbonat-o ieae of insufficien toi producethvolumedo t fuctoncrse-d crystalsfor continued carbonate formation. The preservation field of organic soft parts by carbonatemineralsis therefore controlled by a high input of organic carbon. Rapid burial of such organic-rich sediments would promote an anaerobic regime, reducingthe opportunity bioturbation for and inhibiting iron and sulfate reduction by removing decaying carbon fromthe overlying waterbody. More importantly, residencetime at the anoxic/Ioxic interfaceis reduced, and

KONSERVAT-LAGERSTATTEN

341

the precipitationswitch is leftat the default position (i.e., carbonate on; phosphate off) otherthan as (Fig. 3). Salinityhas littleeffect a control on the species of carbonate which can form.The field of exceptional preservation for carbonate mineralization (Fig. 5) is therefore controlled by a high rate of burial and a high organic content. The Mazon Creek biota is preservedin sideriteconcretionsin the Upper Carboniferous FrancisCreek Shale (Richardsonand Johnson 1971; Baird 1979; Baird et al. 1985). This shale is a greycrevasse-splay deposit(Shabica 1979), which formspart of a cyclothemicsequence. The shale is organic-poor, and the fossil-bearing concretionsare restricted the lowest 10 to m overlying the Colchester (Number Two) Coal. The bicarbonate ions required for sideriteformation originatedfromanaerobic dethis compositionofthepeats which now form coal. A scattereddistribution organic matof ter promotes this type of preservation.A reduced number of precipitation nuclei coupled with increased supply of bicarbonate increasesthe rateofprecipitation and reduces loss. decay-induced information Fine-grained, thinly bedded limestones, such as the JurassicSolnhofen Limestone of Germany,the Eocene Green River Formation of the central U.S. (Grande 1984), and the Haquel and Hjoula Limestones of Lebanon (Huckel 1970; Hemleben 1977), are oftenreto ferred as Plattenkalk deposits or lithographic limestones. Such deposits occur in lacustrine as well as marine systems where the supply of terrigenoussediment is limited.In such a system, respiratory carbondioxide produced by bacteria, algae, and plankton achieves high levels of supersaturationand promotesthe precipitationof carbonateminerals (for review see Bathurst 1975). Rapid burial in such fine-grainedcarbonates promotes the formation of exceptionally preserved fossils. Further,organic productivity may lead to the development of anoxic conditions which inhibit bioturbationand scavenging. Fossils are characteristically flattened and eithercastby the carbonatesor coated by other minerals. There has as yet been little work on their paragenesis or preservational mineralogy.

Conclusion decay inhibitionon a geological Long-term time-scale is an unusual event and largely biological structures restrictedto refractory and unique depositional systems(peat bogs, amber, tarsands,salt deposits, etc.). In more common depositional settings, organic remains can be rapidly destroyedby respiring microbes.Anoxia retardsdecay but does not organic molecules have difhalt it. Different ferentdecay rates; tissues such as muscle are extremely volatile and can be destroyedwithin days or weeks of burial (Allison 1988a). Some compounds, however, are more decayresistant and thereforehave a higher residence time in the sediment.Thus, chitinwill persist in sediment longer than muscle (Allison 1988a), although cellulose will persist for even longer (Allison 1987), and lignified cellulose foreven longer still (Leo and Barghoorn 1970; Stout et al. 1981). According to of Stout et al. (1981), the half-life cellulose is about one-thirdthatof lignin in aerobic conditions. By comparing the relative decay resistance of the tissueswhich formmostof the fossilsthatpaleontologistsencounter,we can see that fossil remains at the outcrop represent the end-product of the interplay between decay and diagenetic mineral formation. If decay progresses aerobically, then diagenetic mineral formationwill be hindered, and organic tissues will be largely destroyed.In such an instance,the sole remaining fossilswill be hard partssuch as bone and shell. Should decay progress anaerobically, then decay rate will be reduced and the conditions required forearly diagenetic mineral formationwill be enhanced. Were mineralization to occur shortlyafterburial,then volatile soft tissues may be preserved. If, h.lowever,mineralizationoccurslaterin the burial historyof the sediment,then only the more tissuessuch as chitin,cellulose,and refractory lignin will be preserved.In the extremecase, tissues are not mineralized at all; then chitin tissue will be destroyed, highlyrefractory but possuch as lignin may remain.It is therefore sible to createa list of biological tissuesbased upon comparative decay resistance (see Fig. 6). This list can be related to the formof pres-

342

PETER A. ALLISON

l'

ff

~~~~~~SHELLY FOSSILS|

INERALIZED
_ UCLE
\\\

EARLY

LATE

MINERALIZATION
of 6. The relationshipbetween decay and mineralizationin the preservation exceptionallypreservedfossils. preservation.If mineralization Reduced decay and very early diagenetic mineralizationare required forsoft-tissue elementsofa biotawill be preserved.The preservational is impeded, decay will continue,and only themorerefractory mineralogyof each tissue type extends fromthe bottomleft-handcorner of the box to the boundary fence of the next most refractory tissue type.
FIGURE

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Acknowledgments J. D. E. G. Briggs,V. P. Wright, D. Hudson, E. N. K. Clarkson commentedon an earand I of lier draft the manuscript. am also grateful to the editor of this journal and two anonymous reviewersfor"redpenning" a previous version and providing useful and construcThis work was initiatedduring tive criticism. the tenure of Natural Environmental ResearchCouncil Quota Award no. GT4/84/GS/ 59 at the Universityof Bristol,England, and completed during a Friday Harbor Postdoctoral Fellowship at the Universityof Washington.

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