Fossils

Mike Viney

Hermanophyton Taylorii Permineralized Seed Fern

The word fossil is derived from the Latin fossilis, which means "du u!"" #nitially, the term fossil a!!lied to any stran e or interestin material found within rock whether or not it was of or anic ori in $Prothero, %&&', !" ()" Most modern definitions include the conce!t that fossils are evidence of ancient or anisms, which have *ecome a !art of the +arth,s crust" The word ancient is ar*itrary" To some, ancient a!!lies only to e-tinct or anisms while, to others, it im!lies time limits" .rimaldi and +n el $%&&() !oint out many would like to restrict the term fossil to s!ecies that have *ecome naturally e-tinct" They ar ue that havin this knowled e is !ro*lematic" .rimaldi and +n el su est the followin !ractical definition, """"a fossil is the remains or workin s of any s!ecies, livin or e-tinct, that have *een naturally !reserved for several thousand years or more $!" /%)" 0 more common time limit defines fossils as *ein !rehistoric thus1 fossils !reserve remains or activities of ancient or anisms older than 1&,&&& years $.arcia 2 Miller, 1334, !" 1'1 Scho!f, 135(, !" %5)" The ma6ority of fossils are found in sedimentary rocks" 7r anisms *ecome tra!!ed within sediment layers due to the action of water, wind or ravity" Fossils can sometimes *e found in metamor!hic rocks formed from fossiliferous sedimentary rocks altered *y heat and !ressure" Fossils can even *e found in i neous rock created from lahars or !yroclastic flows that entom* trees or other or anisms" Two ma6or ty!es of fossils are reco nized" 8ody fossils reveal the structure of an or anism, while trace fossils reveal the activities of or anisms" There are many reasons to study fossils" The fossil record indicates that different life forms have e-isted at different times revealin the evolution of life on +arth" Fossils and rock ty!es serve as clues to determinin ancient environments" Finally, fossils are the most !ractical way of tellin time in eolo y $Prothero, %&&', !" vii)" 9ow do fossils form and how are they classified into different !reservation ty!es: Taphonomy 0 Fossil re!resents evidence of !ast life that is found in the +arth,s crust" Ta!honomy $"laws of *urial") is the term used to descri*e the !rocess that results in the formation of a fossil" Ta!honomy or the transition of an or anism or !art of an or anism from the *ios!here to the lithos!here is accom!lished in *asically two ste!s" The !rocess of *urial or entom*ment is referred to as *iostratinomy" 0fter *urial or entom*ment, dia enesis *e ins1 the conversion of sediments or other de!osits to rock" 8iostratinomic and dia enetic !rocesses destroy most traces of or anisms" The e-tent of !reservation de!ends u!on what ha!!ens durin *iostratinomy and dia ensis" The *iostratinomy and dia ensis associated with a fossil can reveal much a*out the environment in which the or anism lived" #n other words, how fossils form can often !rovide clues to !ast environments" Lagersttten Fossilization often occurs as a result of ra!id *urial, usually *y water;*orne sediment, followed *y chemical alteration" <a!id *urial and s!ecific chemical environments hel! to

reduce decom!osition from *acteria and fun i" =ecom!osition, erosion, de!osition and rock formation are !rocesses that often destroy soft tissue, so it is the hard !arts of or anisms such as shells, *ones and teeth, which are most often !reserved" 7ccasionally conditions e-ist that allow for !reservation of or anisms in environments that rarely !roduce fossils" +-ce!tional conditions may also hel! to !reserve soft tissue or im!ression of soft *ody !arts" Fossil de!osits with soft;*odied or anisms well !reserved or with terrestrial animals, such as dinosaurs in the Morrison formation are termed La erst>tten" La erst>tten is a .erman word used in minin that denotes a !articularly rich seam and has *een ado!ted *y !aleontolo ist to si nify these rare fossil de!osits *ecause they ive us a window into !ast environments seldom !reserved in the fossil record $Selden 2 ?udds, %&&', !" 5)" Two ty!es of fossil la erst>tten are reco nized" =e!osits that contain vast num*ers of fossils re!resent @oncentration La erst>tten" The !reservation may not *e e-ce!tional, *ut the reat num*ers can *e very informative" @onservation La erst>tten contain fossils with soft *ody !reservation, im!ressions of soft tissue or fossils of well;articulated skeletons without soft tissue !reservation" @onservation La erst>tten are !articularly im!ortant *ecause they !rovide knowled e of soft;*odied or anisms, allow !aleontolo ists to reconstruct !aleoecosystems, and ive insi hts into the mor!holo y and !hylo entic relationshi!s of or anisms $?udds 2 Selden, %&&4, !!" 4;3)" Preservation Types The science of ta!honomy e-!lores the environmental conditions that !romote fossilization" 8oth *ody and trace fossils can form under a variety of circumstances re!resentin multi!le modes of !reservation" @om!are several *ooks on fossils off any li*rary or *ookstore shelf and you will soon realize there is no standard for cate orizin ty!es of fossil !reservation" #n this !a!er we will *riefly descri*e modes of !reservation that one is likely to find in *oth scientific !a!ers and !o!ular *ooks" Molds & Casts

@asts 2 Molds of Trilo*ites Ellipsocephalus hoffi

7r anisms *uried in sediment may decay or dissolve away leavin a cavity or mold" #f the s!ace is su*seAuently filled with sediment, an e-ternal cast can *e made" Molds and casts are three dimensional and !reserve the surface contours of the or anism" 0 mold B

!reserves a ne ative im!rint of the surface, while a cast !reserves the e-ternal form of the or anism $Taylor, Taylor 2 Crin s, %&&3, !" %%)" Sometimes a shell can *e filled with minerals and then dissolve away" The internal cast that remains is termed a steinkern, which in .erman means "stone cast" $Pothero, 1334, !" 5)" Steinkerns are most often re!resented *y the internal molds of mollusks" The !ith cast of Calamites is the most common !lant steinkern" 0s a Calamites tree matured the

Calamites Pith @ast $Steinkern)

center of the stem $!ith) *ecame hollow, develo!in into a tu*e;sha!ed air cavity" The !ith cast !reserves an im!ression of the !ith cavities outside surface, which re!resents the inside vascular and corte- tissue $Taylor, Taylor 2 Crin , %&&3, !" %B)" Most molds and casts do not contain the actual remains of an or anism" Shells, *one, and wood often form as molds or casts" Some trace fossils $ichnofossils), such as tracks and *urrows can form as casts or molds" Tracks and *urrows can !rovide clues to the *ehavior and *iomechanics of an or anism while it was alive" @oncretions often enca!sulate a fossil mold and cast" The siderite $iron car*onate) nodules of Mazon @reek, #llinois !reserve @ar*oniferous a ed animals and !lants as molds and casts $?udds 2 Selden, %&&4, !" 1%&)" #t should *e noted that some authors classify the fossils in Mazon @reek nodules as im!ressions $Danssen, 1353, !" %'1 <ich, <ich, Fenton 2 Fenton, 133/ !!" ';/)" Limestone concretions in Ft" @ollins, @olorado contain the molds and casts of @retaceous a ed mollusks" 0s kids, my friends and # collected multi!le Inoceramus clams from a B;foot diameter concretion" These fossils are found as molds and casts with the cast fillin the mold" #t is im!ortant to realize that many fossil s!ecimens re!resent more than one mode of !reservation" For e-am!le, the cast in a Mazon @reek nodule also re!resents fossilization *y authi enic mineralization, a ty!e of re!lacement" Su*tle !9 chan es created *y the decayin *ody of the *uried or anism caused availa*le iron car*onate to !reci!itate" Thus, the or anism *ecame its own nucleation site for the formation of a siderite nodule" Some of the shells found in the limestone nodules of Ft" @ollins retain altered shell material, which re!resents recrystallization" 0ra onite $calcium car*onate) in the shell recrystallized to calcite, a more sta*le form of calcium car*onate"

'

Ee usually think of casts and molds as e-hi*itin o*vious three;dimensional character" The thin leaves of !lants or the win s of insects can !roduce shallow casts and molds" Shallow casts and molds may take the form of im!rints, or im!ressions, !reservin the three dimensional character of an or anism" #m!ressions of win s are a common insect fossil" +ven some of the relief, like !leatin on the win s can *e !reserved" This is im!ortant *ecause the veins on a win can *e used to key an insect to the family level $.rimaldi, =" 2 +n el, M"S", %&&(, !! '%;'()" #m!rints, im!ressions and many trace fossils, such as, *urrows, insect alleries, and tracks, may re!resent ty!es of molds or casts if they retain their three;dimensional character" Molds and casts are im!ortant *ecause they can faithfully re!licate the e-ternal form of an or anism in a three;dimensional fashion, ivin !aleontolo ist information a*out surface anatomy" Impressions, Compressions & Carbonization Ehen or anisms *ecome tra!!ed and sAueezed *etween sediments they may form com!ressions" Lar er or anisms can *e distorted *y com!ression" 9owever, ood fossils of fish, leaves and insects are often formed *y com!ression"

@ar*onized #nsects #nsect @om!ressions

Fish, #nsect and !lant com!ressions retain or anic material" The or anic matter makin u! the *ody of the or anism may *e altered durin decay and rock formation" The distillation of volatile com!ounds and the !olymerization of li!ids chemically transform the or anic structures and leave a thin film" The thin, dark, film is made of sta*le, !olymerized car*on molecules that remain after more volatile and unsta*le com!ounds et dissolved away, hence the name car*onization" 0 com!ression fossil re!resents a ty!e of car*onization" The thin car*on layer on a !lant com!ression is known as a !hytoleim $@leal 2 Thomas, %&&3, !" ')" The !hytoleim may retain ori inal cuticle, which resists decay" The cuticle is the !rotective non;cellular wa-y coverin of the e!idermis" Ehen removed and studied the cuticle may reveal the arran ement of e!idermal cells and stomata, which can sometimes aid in s!ecies identification $Tidwell, 1334, !" %5)"

0s or anisms are sAueezed into com!ressions they may form an im!rint or im!ression" Thus, fossils discovered *y s!littin *eddin !lanes may reveal two fossils from a sin le s!ecimen" The side with more or anic material is called a com!ression" @om!ressions often show the e-ternal surface of an or anism flattened in a two dimensional fashion" The side with little or no or anic material is called an im!ression $Tidwell, 1334, !" %51 Taylor, Taylor 2 Crin s, %&&3, !" %11 Scho!f, 135(, !" B5)" #m!ressions often re!resent a ne ative im!rint of a com!ression"

#m!rint 2 @om!ression of Fish Knightia eocaena

#m!rints are shallow e-ternal molds or voids left *y animal or !lant tissue" Ehen the siltstone !ictured a*ove was s!lit into two sla*s the or anic matter adhered to one side" The to! !icture re!resents an im!rint in which *ones and scales left a shallow e-ternal mold" The lower !icture is a com!ression *ecause it !ossess or anic residue left from scales, ori inal *one, and *one reinforced with calcite" Fish, leaves and insects are often found as im!rints and com!ressions" The com!ression is referred to as the !art $!ositive side) and the im!ression as the counter!art $ne ative side)" The im!ression in this case shows all the surface details of the com!ression and re!resents an im!rint $Taylor, Taylor 2 Crin s, %&&3, !" %1)" The counter!arts of .reen <iver fish that re!resent im!rints can *e used to make !ositive layte- casts to study e-ternal surface features $.rande, 134', !!" 113 2 1%&)" #f the layer of car*on is lost on the com!ression throu h weathernin or further dia enesis then it is

also known as an im!ression $@leal 2 Thomas, %&&3, !" ')" Thus, an im!ression, unlike an im!rint, can *e !ositive or ne ative" For the !aleontolo ist that studies insects, im!ressions are like casts and molds, which may !reserve some relief like !leatin on win s $.rimaldi 2 +n el, %&&(, !" 'B)" This is im!ortant *ecause win venation can *e used to identify an insect" Many Mazon @reek nodules do not retain or anic material and so *oth the !art and counter!art are referred to as im!ressions $Danssen, 1353, !" %'1 <ich, <ich, Fenton 2 Fenton, 133/ !!" ';/)" Sometimes !arts of a s!ecimen are !reserved as a com!ression while other !arts are an im!ression" #n this case the term ad!ression may *e used" 0d!ressions form when the matri- of a fossil is soft and the !hytoleim has fallen off in some !laces $@leal 2 Thomas, %&&3, !" ')" #nsects and leaves !reserved in the +ocene a ed Florissant *eds of @olorado are often car*onized" #t is *elieved that leaves and insects *ecame entan led in diatom mucus mats $formed *y a re ates of diatoms under stress)" #nsects and leaves were incor!orated into layers of sediments and volcanic ash at the *ottom of Lake Florissant" Many of these insects and leaves decom!osed leavin im!rints" 0s the sediments com!acted and hardened into shale the im!rints *ecame im!ression fossils" Some or anisms only !artially decayed retainin a dark colored car*on residue to *ecome com!ression fossils $car*onization)" Many insects have their win s !reserved as im!ressions and their *odies as dark com!ressions" @om!ressions are often flattened, havin a two;dimensional a!!earance" 9owever, the !reservation in diatom layers allows some or anisms to retain their three;dimensional character" Some insects are found with or ans and a!!enda es" Some leaves can *e found with internal structures $Meyer, %&&B, !!" B(;B5)"

@one 2 ?eedle @om!ression of Metasequoia

Feathers are often !reserved throu h com!ression and car*onization" #t is *elieved that the car*on residue is the result of feather de radin *acteria" 0n analysis of a @retaceous a ed fossil feather showin a *anded color !attern from 8razil !roduced interestin results" The li ht areas of the feather re!resented an im!ression $no or anic residue)" The dark areas, re!resentin a com!ression, consisted of 1;% micrometer o*late car*onaceous *odies" These o*6ects turned out to *e car*onized melanosomes1 molecules that were 5

res!onsi*le for the ori inal feather color" The fact that the structure of the melanosomes is !reserved o!ens u! the !ossi*ility of determinin the ori inal feather color $Vinther, D", %&&4, !" (%%)"

Fusainized Lithocarpus sp.

Most fossils found in coal de!osits are com!ressed and car*onized or coalified" @oal *alls are the e-ce!tion and are discussed under !ermineralization" Plant and animal remains may also *e @harcoalified or Fusainized" Many *elieve that fusainized or anisms are transformed to charcoal *y ancient forest fires" 0lthou h evidence of fire is associated with some fusainized !lant tissues there are e-ce!tions" Some fusainized remains !reserve cuticle and resins, which does not seem consistent with an ori in that includes fire $Scho!f, 135(, !" '()" Fusainized remains are three;dimensional and may *e re!licated to the cell level *y car*on $.rimaldi 2 +n el, %&&(, !!" '3;(&)" Permineralization

Permineralized 7ak Quercus sp.

Permineralized fossils form when solutions rich in minerals !ermeate !orous tissue, such as *one or wood" Minerals !reci!itate out of solution and fill the !ores and em!ty s!aces" Some of the ori inal or anic material remains, *ut is now em*edded in a mineral matri4

$Scho!f, 135(, !" B1)" 8one and wood tissues act as e-cellent frameworks to !reserve cell structure" Silicates, iron o-ides, metal sulfides, native elements, car*onates, and sulfates can *e involved in !ermineralization" Permineralization is one of the most faithful modes of fossil !reservation" #n fact, scientists have tried to re!licate the !rocess in the la*oratory, *ut no artificial !ermineralization is eAual to the *est natural !reservation *y cry!tocrystalline silica or calcium car*onate $Scho!f, 135(, !" BB)" Formation of the finest !etrified wood involves !ermineralization with silica, usually from a volcanic source, alon with re!lacement and recrystallization" =urin the initial sta es of !ermineralization amor!hous silica infills !its connectin cells and !reci!itates on cell walls" 0t this early sta e no re!lacement has occurred" <e!lacement of cellulose in cell walls may occur as !ermineralization continues" @ellulose that de rades leaves room for the em!lacement of silica *etween and within cells walls" The more decay resistant li nin that remains in the cell walls continues to act as a uidin framework to !reserve structure" Later silica is de!osited in cell lumina, the cavity enclosed *y the cell walls and voids left *y wood de radation" Silica that initially !ermeates the !orous tissue and that which re!laces cell wall material is amor!hous" This amor!hous silica is unsta*le and slowly crystallizes to more sta*le forms over millions of years" The transition to more sta*le forms of silica involves continued !olymerization and water loss" 9i her ordered forms of o!al are created throu h this !rocess and eventually lead to the thermodynamically more sta*le silica Auartz $Stein, 134%, !" 1%55)" The Auality of !reservation usually, *ut not always, declines durin successive sta es of silicification $Mustoe, %&&B, !" B/)" #n some instances hi her order forms of o!al and chalcedony may act as the initial re!licatin minerals $Mustoe, %&&4, !" 1B4)"

Permineralized Palmoxylon

Petrified forests, re!resentin small to lar e de!osits of !ermineralized wood, ca!ture !eo!le,s ima ination" Ehat !rocesses allow wood structure to *e !reserved in stone: 3

9ow lon does it take to form !etrified wood: Fou can e-!lore these Auestions in more de!th *y readin our article on Permineralization" Petrified wood and !etrified dinosaur *one are !ro*a*ly the *est known !ermineralized fossils amon the eneral !u*lic" 0lthou h not as well known, the coal *all re!resents a very informative !ermineralized fossil" 0 s!ecial ty!e of fossil, the coal *all, can *e found in the coal de!osits of the Pennsylvanian and Permian !eriods" @oal *alls contain swam! ve etation, which has *een !ermineralized with calcium car*onate, !reservin B; = cellular structure" @oal *alls are studied in serial section usin the cellulose acetate !eel method to reveal microsco!ic structure" Serial sections can *e used to reconstruct or ans and entire !lants" The five ma6or rou!s of !lants found in coal *alls includeG Lyco!hytes, s!heno!sids, ferns, seed ferns, and cordaiteans $<othwell, %&&%, !" '&)" The in situ !reservation of !lant materials in coal *alls allows !aleontolo ist to study !lant associations that tell us somethin a*out the !alaeoecolo y of the coal swam!s" @oal *alls reveal that the ar*orous fern Psaronius *ecame the dominant cano!y tree after the e-tinction of Le!idondendrales near the Middle Pennsylvanian" @ertain s!ecies of small ferns and horsetails have *een found, which rew in association with the roots of Psaronius $<othwell, %&&%, ! '%)" Amber

Eas! in 8altic 0m*er

0m*er is referred to as !etrified tree resin or sa!" # !refer !etrified tree resin as the term sa! refers to fluids trans!orted *y -ylem or !hloem tissues $<aven, +vert, 2 @urtis, 1341, !" /(3)" @onifers and some deciduous trees !roduce resin in res!onse to in6ury" <esins are viscous liAuids that contain volatile ter!ene com!ounds and or anic solids" Hnder the ri ht conditions resins !olymerize and harden with a e, turnin into co!al" 0fter several million years or more, co!al matures into am*er" Tree resin *reaks down when e-!osed to dryin and o-idation within 6ust a few thousand years" #t is not sur!risin then that am*er de!osits do not re!resent forest floor environments" 0m*er de!osits usually re!resent marine environments" 0m*er de!osits form when resins !roduced in forests are trans!orted *y water to oceans or lakes, where they are de!osited into the sedimentary layers" Iuick trans!ort and de!osition !rotects 1&

the resin from weatherin " 7nce de!osited, the resin chemically matures into intermediate forms called co!als and finally into am*er after millions of years" The am*erization !rocess is estimated to take *etween % and 1& million years" 9owever, the ty!e of de!ositional environment may also affect the time needed for am*erization" 0m*er from 8orneo is found in sand and clay sediments de!osited in a dee! ocean 1% million years a o" The material that comes out of the sandstone has matured into am*er, while the s!ecimens from the clay are still co!al $<oss, %&1&, !! 4;3)" Petrified resins have *een found in @ar*oniferous, Triassic, and Durassic de!osits, *ut re!resent minute amounts of resins !roduced inside trees" <esin that collects inside trees does not act as an insect tra!" The first occurrence of fossil containin am*er is @retaceous in a e" The ma6ority of am*er de!osits that contain fossils were formed durin the @enozoic $Eeitschat 2 Eichard, %&&%, !!"3;1&)" Fossils entom*ed in am*er are referred to as inclusions" 0lthou h the or anisms often look com!lete, most a!!ear to *e thinly lined hollow s!aces $Eeitschat 2 Eichard, %&&%, !" %3)" 9owever, under the ri ht conditions the internal or ans can *e well !reserved" The !reserved internal or ans of a *ee e-humed from =ominican am*er have *een ima ed usin an electron microsco!e $.rimaldi 2 +n el, %&&(, !" (3)" Studies usin scannin electron microsco!es as well as transmission electron microsco!es have revealed internal or an !reservation in 8altic am*er s!iders and nats" #t seems that many or anisms in am*er are !reserved throu h mummification" #n the !rocess of mummification, dehydration results in u! to a B&J decrease in volume of tissues" The decrease in tissue volume ives the or anisms the a!!earance of an em!ty husk $Selden 2 ?udds, %&&', !" 1B')" The sha!e of a s!ecimen can *e a clue to whether the am*er formed inside the tree as an internal resin accumulation or outside the tree as an e-ternal resin accumulation" <esin can collect inside the void of a tree, dri! off a *ranch, or flow alon the outer *ark" <esin that collects inside a tree usually does not contain fossils" <esin that accumulates on the outside of a tree can act as an insect tra!" Fossils are almost e-clusively found in s!ecimens formed *y successive resin flows that collected on the outside of the tree" These s!ecimens are referred to as Schlau*e $Eeitschat 2 Eichard, %&&%, !" 1%)" 7r anisms *ecome tra!!ed in the resin and are then covered *y a successive resin flow" Fou can usually see the !lane re!resentin a successive resin flow1 it often looks like a fracture in the am*er" 0m*er can re!resent a *rief sna!shot in time" 0m*er has !reserved insect develo!mental sta es, matin , e layin , *rood care, feedin , as well as various sym*iotic relationshi!s" 0m*er with and without fossils can *e Auite valua*le so, it is often faked" See <oss $%&1&) for an e-cellent discussion re ardin materials used to make fake am*er and tests that can *e used to distin uish real am*er from imitations $!!" 11;1()"

11

Chemical Preservation

Hy rophilus *eetle in Tar Sand

Peat *o s, oil see!s, !araffin, tar !its and as!halt are ood sources for fossils that have *een chemically !reserved $.arcia 2 Miller, 1334, !" 1()" Paraffin mines and !eat *o s can !reserve soft tissue" Tar !its or as!halt !reserve only hard !arts such as *ones, shells or e-oskeletons" #n 13&5, a !araffin mine in Poland !roduced the head, forele s, and skin of a woolly rhinoceros $<ich, 133/, !" B)" The <ancho La 8rea Tar Pits in @alifornia, 8i 8one Lick in Centucky, and Talara, Peru are well known sites for fossils !reserved in tar !its $.arcia 2 Miller, 1334, !" 1()" <ancho La 8rea literally translates to "the tar ranch"" The naturally occurrin !etroleum *ased su*stance is more !ro!erly referred to as as!halt $?udds 2 Selden, %&&4, !" %/%)" <ancho La 8rea within the city of Los 0n eles is one of the richest de!osits of ice a ed animals" The sa*re;toothed cat !milo on fatalis, the im!erial mammoth Mammuthus imperator, the 0merican mastodon Mammut americanum, and the iant round sloth "lossotherium harlani are 6ust some of the ice a ed mammals that ca!ture the !u*licKs ima ination" 9owever, hundreds of !lant and animal s!ecies have *een found tra!!ed within these as!halt de!osits !rovidin a window into a ?orth 0merican Pleistocene ecosystem" Most of the fossils e-cavated at <ancho La 8rea are car*on dated at *etween 11,&&& and B4,&&& years old" The as!halt rich sediments that contain this concentration la erst>tte were de!osited durin the last ice a e $Eisconsinan .laciation), which !laces them in the H!!er Pleistocene +!och" Shallow as!halt !ools formed animal tra!s durin the summer" =urin the winter these !ools may have solidified" 0s seasons chan ed, heat from the summer sun would once a ain set the tra!s for fora in her*ivores" #nterestin ly, there is a !reservation *ias for carnivores, youn and maimed individuals" Foun and maimed animals were more susce!ti*le to *ecomin tra!!ed in the as!halt !ools" Scaven ers were attracted to the carcasses accumulatin in the !ools" 0lthou h

1%

soft tissue is not !reserved the *ones retain much of their ori inal com!osition" <a!id *urial followed *y as!halt !ermeation accounts for the e-cellent *one !reservation $?udds 2 Selden, %&&4, ! %/%;%/4)" The *ones are *lack with tar and have the smell of !etroleum" Scientists have e-tracted =?0 from the *ones to com!are these e-tinct or anisms with their livin relatives $Prothero, %&&', !" 3)" eplacement

Pyritized 0mmonite

Minerals can re!lace *one, shell, wood, and even soft *ody !arts as they dissolve away due to the action of water and decay" <e!lacement and mineralization are terms used to descri*e this fossilization !rocess $.arcia 2 Miller, 1334, !" 1()" Part of the ammonite shell a*ove has *een re!laced *y the mineral !yrite" The re!lacement of soft or hard *ody !arts may occur when minerals !reci!itate out of solution due to the action of *acteria or !9 chan es" =urin re!lacement coats of *acteria Auickly mineralize the decayin tissue" #f re!lacement results in a fossil that is com!letely articulated with three;dimensional fidelity the !rocess is referred to as mineral re!lication $.rimaldi and +n el, %&&(, !" '()" .rimaldi and +n el also classify !ermineralization as a ty!e of mineral re!lication that is a result of micro*ial decay" 7r anic residue on com!ression fossils can *e re!laced *y minerals leavin an im!ression coated with a mineral" Pyrite is a common re!lacement mineral" #n !yritization sulfur reducin *acteria facilitate the !reci!itation of !yrite durin decay" The La erst>tten known as 8eecherKs Trilo*ite 8ed in ?ew Fork is famous for its !yritized trilo*ites" Trilo*ite e-oskeletons as well as soft *ody !arts, includin antennae, le s, muscles, and di estive tract, have *een !reserved with the mineral !yrite $+tter, %&&%, !" 1B1)" #n the 7rsten de!osits of Sweden the meiofauna makin u! the !aleoecosystem of a flocculent;layer 6ust a*ove the sea*ed is e-Auisitely !reserved *y !hos!hatization" +yes, hairs, s!ines, muscle scars, 6oints, !ores, and soft *ody !arts have *een !reserved on miniature late @am*rian arthro!ods" +-oskeletal re!lacement and or coatin *y calcium

1B

!hos!hate occur only on s!ecimens less than % mm is size" The 7rsten La erst>tten is im!ortant *ecause it hel!s to dee!en our understandin of arthro!ods as many larval sta es are !reserved $Tan , %&&%, !!" 115;1%1)" +-Auisite e-am!les of leaves, stems, cones, and seeds of @ar*oniferous !lants alon with animal life can *e found in the La erst>tten known as Mazon @reek, which is 6ust 1(& km southwest of @hica o, #llinois" The soft and hard !arts of !lants and animals are re!laced with the mineral siderite $iron car*onate)" Su*tle !9 chan es created *y the decayin *ody of the *uried or anism caused availa*le iron car*onate to !reci!itate" Thus, the or anism *ecame its own nucleation site for the formation of a siderite nodule" Ehen these nodules are s!lit o!en, the fossil a!!ears as a B;= e-ternal cast and mold" Mold surfaces may *e coated with kaolinite, !yrite, calcite or s!halerite" Plant material is sometimes covered with a car*onaceous film $?udds 2 Selden, %&&', !" 1%&)"

Siderite ?odule with Seed Fern Leaf

Plant material !reserved in *arite sand nodules can *e found near the town of Steinhardt .ermany in an 7li ocene a ed de!osit" Ehen s!lit o!en some of these nodules reveal molds and casts of !lant material re!laced with the mineral *arite $*arium sulfate)"

1'

@one @ast in 8arite ?odule

The !rocess that forms a concretion or nodule, which may contain a re!lacement fossil, is called authi enic cementation or authi enic mineralization $Prothero, %&&', !" 'B51 @leal 2 Thomas, %&&3, !" 5)" 0uthi enic minerals row in !lace rather than *ein trans!orted or de!osited" Thus, the concretions or nodules are found in the !lace they formed or in situ $Latin for "in the !lace")" ecrystallization

<ecrystallized Fossil @lam Mercenaria permagna

Some shells are made of ara onite" =urin the fossilization !rocess ara onite reverts to a more sta*le form of calcium car*onate called calcite" Thus, recrystallization from 1(

ara onite to calcite re!resents a ty!e of re!lacement" Some shells are made of layers of calcite and ara onite" The small crystals of calcite in shells may recrystallize into lar er calcite crystals" The overall sha!e of the shell may remain, *ut the effect of recrystallization on microsco!ic te-ture is evident $Prothero, %&&', !" 3)" Freezing

Mammuthus primigenius 9air

Eoolly mammoths and woolly rhinoceroses from the Pleistocene can sometimes *e found in the !ermafrost of 0laska and ?orthern Si*eria" The entire or anism is sometimes !reserved in this frozen soil" =urin the last #ce 0 e some of these animals died in areas that have remained cold" +ventually, in rare instances, or anisms were *uried in what *ecame !ermafrost soil" 8ones, muscles, internal or ans, !artly di ested food, skin and hair can sometimes *e found" 8oth the course uard hair and soft underwool of the mammoth is re!resented in the s!ecimen a*ove" Some of the B&,&&& year old woolly mammoths found are so fresh that they could *e eaten *y humans and animals $Prothero, %&&', !" 3)" #n the s!rin of %&&5 Furi Chudi, a ?enet reindeer herder, discovered a *a*y Mammuthus primigenius e-!osed on a sand*ar of the Furi*ey <iver in Si*eria" The '&,&&& year old fossil mammoth was named Lyu*a after ChudiKs wife" Lyu*a, nicknamed the #ce 8a*y, re!resents one of the *est !reserved fossils found to date in the !ermafrost of Si*eria" Lyu*a was one month old when she drowned in soft sediments of silt and clay" Paleontolo ist =an Fisher has determined that more than 6ust the frozen !ermafrost was essential in Lyu*aKs e-cellent !reservation $Miller, %&&3, !" '1)" Lacto*acilli colonized her tissues after death" The lactic acid !roduced *y these *acteria acted as a !reservative, !icklin Lyu*aKs tissues" 0s new sediments accumulated a*ove 1/

Lyu*a the layers in which she was entom*ed turned to !ermafrost" +ventually, floodwaters eroded the !ermafrost that encased Lyu*a and trans!orted her downstream" The lactic acid that ori inally hel!ed to !reserve the tissues now !rotected the fossil from !resent day scaven ers as it lay e-!osed on the sand*ar" Thus, for Lyu*a !reservation *y *oth chemical means and freezin were critical factors in the fossilization !rocess" !ncaps"lation +nca!sulation occurs when minerals form around an or anism" Late Miocene y!sum de!osits in the 0l*a area in Piedmont, ?orthern #taly contain dra onflies $mostly larvae) entom*ed in sin le clear y!sum crystals" The insects *ecame tra!!ed in the y!sum as the eva!orate de!osit formed" The entom*ed insects, like most inclusions in am*er, are thinly lined hollow s!aces $Schluter, Cohrin , 2 .re or, %&&B, !" B5')" 7r anisms may also *ecome entom*ed in microcrystalline material" The =evonian a ed <hine @hert near the 0*erdeenshire villia e of <hynie in Scotland re!resents an ecosystem near a sinter terrace" The area was !eriodically flooded with silica rich solution from hot s!rin s and eysers $Selden 2 ?udds, %&&', !" (% and Cenrick 2 =avis, %&&', !" %')" 7r anisms were !ermeated with silica and entom*ed *efore any cellular decay could occur" #nsects have also *een found enca!sulated in Miocene a ed ony- from 0rizona $.rimaldi 2 +n el, %&&(, !!" '3;(&)" #esiccation =esiccation occurs when an animal dies in a very dry environment" Eater is drawn out of the tissues slowin the !rocess of decay" The dryin !rocess may also reduce the !ro*a*ility of scaven in " This !rocess is similar to human mummification" #n fact, some authors use the term mummification to descri*e this !rocess" .round sloths !reserved throu h desiccation have *een found in South 0merica $.arcia 2 Miller, 1334, !" 1()" Moa remains !reserved throu h desiccation have *een found in ?ew Lealand $Ealker 2 Eard, %&&%, !" 1B)" Many of these s!ecimens have *een found in dry caves" ?aturally mummified insects have *een found in association with Pleistocene mammals frozen in tundra !ermafrost" #nsects !reserved throu h desiccation have also *een found in + y!tian mummies and the stomachs of +ocene a ed *ats $Martinez;=elclos 2 Darzem*owski, %&&&)" Ehen is a !reserved remain considered a fossil: Ealker and Eard $%&&%) do not consider or anisms !reserved throu h desiccation to *e fossils *ecause they are only tem!orarily s!ared from decay $!" 1B)" +arlier we discussed that some limit the definition of fossils to the remains of a s!ecies that have *ecome naturally e-tinct" Many definitions have time limits, which are set somewhat ar*itrarily" .imaldi and +n el $%&&() su est that evidence of any or anism which has *een naturally !reserved for several thousand years or more constitutes a fossil $!"/%)" Hnder this definition desiccation may *e considered a fossilization !rocess"

15

0s discussed earlier, desiccation also seems to !lay a role in the !reservation of some or anisms enca!sulated in am*er" Tree resin !rovides a micro;environment in which mummification can take !lace" 0s the resin matures into am*er the mummified remains continue to *e !rotected from decay" $naltered emains

Mammuthus primigenius molar

The conce!t of unaltered remains can refer to multi!le modes of !reservation" Freezin , enca!sulation in am*er $tree resin), desiccation, and chemical !reservation, such as entom*ment in !etroleum containin sediment, are e-am!les e-!lored in our museum" The term unaltered remains is a *it misleadin " #t does not mean that the or anism is unchan ed" ?ucleic acids $=?0 and <?0), !roteins, !i ments, and soft tissues may *e de raded" Tissues, if !resent, have usually lost water" 9owever, or anic matter that is !resent has not chan ed into another su*stance" Freezin , mummification $desiccation), oil see!s, and am*er can !reserve *oth soft and hard tissues" Sometimes the soft tissues decay, *ut the hard !arts remain unaltered" Teeth, *ones, and shells may *e !reserved in this way" Forty thousand year old *ones encased in the as!halt of <ancho La 8rea in Los 0n eles retain their ori inal com!osition" Mollusk shells of the Pleistocene are known that retain their mother;of;!earl ara onite layer" +ven some @retaceous a ed mollusks are found with their ara onite intact $Prothero, %&&', !" 3)" The mammoth tooth !ictured on this !a e retains ori inal enamel, dentine, and *one" Hnaltered teeth, *ones, and shells may also e-hi*it some si ns of !ermineralization with calcium !hos!hate, !yrite or other minerals"

14

Chemical Fossils

@rude 7il

@hemical fossils are chemicals found in rocks that !rovide an or anic si nature for ancient life" Molecular fossils and isoto!e ratios re!resent two ty!es of chemical fossils" Molecular fossils are often referred to as *iomarkers or *iosi natures and re!resent !roducts of cellular *iosynthesis that are incor!orated into sediments and eventually into rock" Many of these chemicals *ecome altered in known ways and can *e sta*le for *illions of years" ?ucleic acids $=?0 2 <?0), !roteins, and car*ohydrates do not survive lon in the eolo ic environment" The ma6ority of *iomarkers are hydrocar*ons derived from mem*rane li!ids, which under certain conditions can *e sta*le over *illions of years" Molecules derived from !i ments, such as chloro!hyll, can also act as *iomarkers" #n 13B/ 0lfred Tei*s reco nized that vanadyl !or!hyrin was a molecular fossil of chloro!hyll" Tei*s discovery hel!ed su!!ort a *iolo ic ori in for !etroleum $Cnoll, summons, Eald*auer, 2 Lum*er e, !!" 1B';1B()" The fossil fuels !etroleum $crude oil), coal, and natural as are the result of *iolo ic activity and contain chemical fossils" Ma6or coal de!osits re!resent !lant material that rew !rimarily durin the @ar*oniferous !eriod" @rude oil and natural as formed !rimarily from !rehistoric al ae and zoo!lankton that were de!osited on the ocean floor under ano-ic conditions" ?atural as can also form from fossil !lant material" =urin sedimentary rock formation the remains of al ae and zoo!lankton are converted into a mi-ture of or anic hydrocar*ons known as kero en" 7ver eolo ic time heat and !ressure can convert kero en into oil or natural as" The ma6ority of oil de!osits are Mesozoic or @enozoic in a e"

13

#t is interestin to contem!late the ori ins of fossil fuel ener y" 0ncient !lants and al ae converted solar ener y into the chemical *ond ener y of car*ohydrates" @onverted ener y from the Sun was then !assed throu h the food chains of these !rehistoric ecosystems" 7r anic chemicals from these or anisms were incor!orated into sediments and eventually rock" So, the fossil fuels we humans have come to de!end u!on re!resent ancient sunshine stored within the +arthKs crust" Fossil fuels currently !rovide more than 4(J of all the ener y consumed in the Hnited States" @rude oil su!!lies '&J of our ener y needs and accounts for 33J of the fuel for cars and trucks" @oal is the ma6or source of ener y for eneratin electricity worldwide $H"S" =e!artment of +ner y)" Ee de!end u!on fossil fuel ener y for our a riculture" Some studies estimate that without fossil fuels the Hnited States could only sustain two thirds of its current !o!ulation $Pfeiffer, ="0", %&&/, !" '1)" 7ur de!endence u!on these nonrenewa*le resources should *e a wake u! call to invest in and develo! alternative ener y sources" 0t !resent, it is clear that coal, oil and natural as, chemical fossils in the form of fossil fuels, are the life*lood of 0mericaKs economy" For !aleontolo ists and eo*iolo ists the information !rovided *y molecular fossils varies reatly" Some molecular fossils can hel! to determine what or anisms were !resent, while others can indicate what *iosynthetic !athways were in o!eration1 still others !rovide information re ardin the de!ositional environment $Cnoll, Summons, Eald*auer, and Lum*er e, %&&5, !" 1B()" #soto!e ratios re!resent another ty!e of chemical fossil and result from meta*olic !rocesses that !referentially utilize one form of an isoto!e over another $@owen, <" !" 1/)" Molecular fossils re!resent *iomolecules or their derivatives that were once !art of a livin or anism" #n this way, molecular fossils are similar in conce!t to conventional *ody fossils" #soto!e ratios are not !reserved *its of an or anism, rather they result from activities durin life and in this way are analo ous to trace fossils" The first chemical evidence of !hotosynthesis, in the form of @;1% to @;1B ratios, can *e found in 0rchean rock of B"4 .a from #sua, .reenland $Cenrick 2 =avis, %&&', !! 1&; 111 Dohnson 2 Stucky, 133(, ! %%)" The !rocess of !hotosynthesis !referentially utilizes @;1% over @;1B when removin @7% from the air to synthesize car*ohydrates, creatin ratios of these car*on isoto!es that differ from normal *ack round ratios" Thus, car*on com!ounds !rocessed *y !hotosynthetic or anisms are enriched with @;1% $<ich 2 Fenton, 133/, ! 31)" The enrichment of @;1% in rocks is a test for the !resence of life" @ar*on isoto!e ratios consistent with !resence of cyano*acteria are wides!read in rock dated at B"( .a $Dohnson 2 Stucky, 133(, !" %%)" 7ne !ro*lem with the evidence a*ove is the fact that chemical !athways in non;!hotosynthetic autotro!hs and nonautotro!hs can !roduce @;1% enrichment $8lankenshi!, Sadekar, 2 <aymond, %&&5, !! %%;%B)" Molecular fossils have *een a key to understandin the evolution of !rimary !roducers in +arthKs oceans" Microfossils and molecular fossils have hel!ed to esta*lish that +arthKs oceans have e-!erienced two ma6or shifts in the com!osition of !rimary !roducers" #nitially, cyano*acteria alon with other !hotosynthetic *acteria were the !rimary

%&

!roducers durin the Proterozoic eon" The first shift occurred durin the early Paleozoic era when eukaryotic reen al ae 6oined cyano*acteria in *ein ma6or !rimary !roducers" The second shift would occur durin the Mesozoic era when dinofla ellates and coccolithofores would *e 6oined *y diatoms in the Durassic" =iatoms, dinofla ellates, and coccolitho!hores would assume their dominant role as the *ase of many modern marine ecosystems *y @retaceous times" $Cnoll, Summons, Eald*auer, and Lum*er e, %&&5, !" 1(()" Trace Fossils

Trilo*ite Tracks

Fossils do not always re!resent a !art of the or anism" Trace fossils record the activities of or anisms" Tracks, *urrows, e shells, nests, tooth marks, astroliths $ izzard stones), and co!rolites $fossil feces) are e-am!les of trace fossils or ichnofossils" Trace fossils re!resent activities that occurred while the animal was alive" Thus, trace fossils can !rovide clues to diet and *ehavior" #chnolo y $ichn "trace or track, ;olo y "the science of") is the study of trace fossils" Trace fossils re!resent multi!le modes of !reservation and are considered here as a cate ory for convenience" Tracks re!resent animals oin a*out their day;to;day activities and may !rovide insi ht into the dynamic *ehavior of e-tinct or anisms" Tracks are formed in situ, that is they are found in the !lace were the or anism made them" The rocks containin tracks !rovide clues to the environment in which the im!rints were made" Foot!rints makin u! a track can reveal the !ace $ste!s), stride $the distance *etween consecutive ste!s made *y the same foot), and trackway width or straddle" Ste!s and stride can reveal anatomical features, such as, num*er of toes or whether the or anism was a *i!ed or Auadru!ed" Straddle can *e used to measure the e-tent to which the animal s!rawls or walks erect $Lockley 2 Meyer, %&&&, !!B;')" Pace an ulation can $an le *etween ste! line se ments) hel! to determine the *ody width of an animal $Prothero, 1334, !" '1B)" %1

Mathematical relationshi!s *etween stride len th and hi! hei ht $measured *y foot!rint len th) of some verte*rates can hel! us to esta*lish relative velocity" Velocity $V) mMs N $&"%(
&"(

)$SL 1"/5)$h ;1"15)

#n 135/ <" Mc?eil 0le-ander, a 8ritish zoolo ist, !ro!osed the most widely used formula for estimatin the s!eed of animals from their trackways where is the acceleration due to ravity, SL the stride len th, and h hi! hei ht $foot!rint len th - ')" <elative stride len th is a ratio of stride len th divided *y hi! hei ht" 0 ratio of reater than %"& for most terrestrial tetra!ods marks the !oint at which an animal chan es from walkin to trottin " 0 ratio of reater than %"3 marks the !oint at which an animal is runnin $Prothero, 1334, !" '1%)" Paleontolo ists have also attem!ted to use tracks as an indicator of meta*olism1 was the or anism endothermic $"warm *looded") or ectothermic $"cold *looded"): Trackways can !rovide clues to the social nature of the animal, was it re arious $social), and did the animal travel in herds: Trace fossils can *e com*ined to !rovide multi!le lines of evidence" =inosaur nestin sites and trackways su!!ort the idea that some her*ivorous dinosaurs were re arious" This same evidence may also !oint to mi ratin *ehavior" Tracks of multi!le or anisms $foot!rint assem*la e or ichnocoenosis) com*ined with an analysis of rock formation can hel! to *uild an ecolo ical !icture of ancient environments" Trackways at =aven!ort <anch in Te-as record a herd of %B sauro!ods a!!arently *ein tracked *y ( thero!od dinosaurs $Prothero, 1334, !" '1') The system for namin foot!rints $ichnota-onomy) runs somewhat !arallel to the ta-onomy for *ody fossils" 0 track made *y Tyrannosaurus would *e iven the formal name Tyrannosauripus" Foot!rint names end in ;!us $"a foot"), ;!odus $"foot"), or ;ichnus $"track or trace") $8orror, 1344, !! '5, 54, 2 4%)" =eterminin who made tracks is a !rime o*6ective of trackin " Lookin at *ody fossils from the same time !eriod to com!are foot anatomy to the track is the key" For e-am!le, Pterosaur feet $*ody fossils) are an e-cellent match for Pteraichnus $foot!rints)" #n very rare instances the tracks are found with the maker" 0 s!ecimen of Kouphichnium #alchi $a horseshoe cra*), found in the Solnhofen strata, is !reserved at the end of its track $8oucot, 133&, !" B1')" 8ivalves with esca!e trails have *een found in siderite nodules from Mazon @reek $?udds 2 Selden, %&&4, !" 1%B)" Patterns of tracks throu h time, known as !alichnostrati ra!hy corres!onds well with *iostrati ra!hic zones $time intervals defined *y fossils)" The lon evity of the avera e dinosaur s!ecies, defined *y a!!earance and disa!!earance from the eolo ic record, is 5 to 4 million years" #chnolo ists find essentially the same s!an of time, 5 to 4 million years for chan es in the foot!rint record $Lockley 2 Meyer, %&&&, !" 1&)" The <everend Eilliam 8uckland $154';14(/), an +n lish eolo istM!aleontolo ist, was the first scientist to reco nize the true nature of fossilized feces" 8uckland coined the term co!rolite or "dun ;stone" to descri*e these trace fossils" The oldest known

%%

verte*rate co!rolites come from Silurian de!osits and re!resent fish feces $+sch*er er, %&&&, @o!rolite 0rticle)" @o!rolites attri*uted to arthro!ods are known from the 7rdovician $Taylor, Taylor 2 Crin s, %&&3, !" 1&&5)"

@rocodile @o!rolite

=r" Caren @hin, curator of !aleontolo y at the Hniversity of @olorado Museum in 8oulder identifies several criteria for co!rolite identification" @o!rolites often have hi h calcium !hos!hate content $Eilliams, %&&4, !" '5)" Phos!hate hel!s mineralize feces" This may hel! to e-!lain why there is a !reservation *ias for carnivore co!rolites over those of her*ivores" @arnivoreKs co!rolites contain their own source of !hos!hate in the *ones and teeth of the consumed !rey" 9er*ivorous co!rolites contain the cellulose and li nin of !lants and reAuire an outside source of !hos!hate, usually in the form of marine sediments" The need for !hos!hate does not a!!ly to all co!rolite !reservation unlike verte*rates, the co!rolites of terrestrial arthro!ods, have *een found in the <hynie chert, !ermineralized coal *alls and fossil wood $Taylor, Taylor, 2 Crin s, %&&3, !!" 1&&5; 1&11)" Sha!e can also *e an im!ortant clue" Sharks and many !rimitive fish !roduce s!iral; sha!ed co!rolites" The sha!e of co!rolites from lar er or anisms such as dinosaurs may *e affected *y s!lattin issues, tram!lin , weatherin and dun consumers" Sha!e can also *e misleadin " Salmon @reek in the state of Eashin ton !roduces *eautifully sha!ed siderite $iron car*onate) de!osits, which resem*le feces" These structures are sold as turtle or even round sloth Miocene;a ed co!rolites" There is some de*ate a*out the nature of these de!osits" 0dolf Seilacher of Fale Hniversity su ests they may re!resent intestinal casts or cololites $Seilacher, %&&1, !" 1)" .eor e Mustoe of Eestern Eashin ton Hniversity !oints out that there is a real lack of evidence as to an animal ori in for these de!osits" First, they lack calcium !hos!hate" Second, they lack internal clues such as !ollen, scales, seeds, *ones or !lant fi*ers" Third, the de!osit in which they are found lacks fossils" These siderite de!osits most likely do not re!resent co!rolites" The real value in co!rolites is what they contain"

%B

Salmon @reek Pseudoco!rolite:

@o!rolite research is carried out !rimarily with thin sections" 9er*ivorous dinosaur co!rolites studied *y @hin have esta*lished a relationshi! *etween !lant eatin dinosaurs and tunnelin insects related to dun *eetles $+sch*er er, %&&&, @o!rolite 0rticle)" =r" @hin has also esta*lished that T$rex !ulverized its victims from studyin microsco!ic *one fra ments found in a co!rolite" #ndian and Swedish scientists have found rass !hytoliths in dinosaur co!rolites" This hel!s to esta*lish that dinosaurs and rasses coe-isted $Eilliams, %&&4, !" (1)" @o!rolites of arthro!ods recovered from rock material, such as the <hynie chert, are com!osed of s!ores as well as !lant and fun al remains $Taylor, Taylor 2 Crin s, %&&3, !" 1&&5)" These =evonian;a ed arthro!ods were !ro*a*ly detritus feeders" @o!rolites !rovide im!ortant insi ht into diet, !hysiolo y as well as the eolo ic and eo ra!hic distri*ution of !lants and animals" #nsect ichnofossils $trace fossils) can *e hel!ful in determinin what ty!es of insects were !resent at a !articular time and !rovide information a*out the nature and !ersistence of !ast !lant;insect associations" +vidence for her*ivory in insects a!!ears in the @ar*oniferous" Like verte*rates, the first insects were carnivores and detritivores" 9er*ivory reAuires hostin cellulose;di estin *acteria throu h a sym*iotic relationshi! within the ut" The oldest e-am!les of mar inal and surface feedin are on @ar*oniferous seed fern leaves of %europteris and "losspteris $.rimaldi 2 +n el, %&&(, !" (%)" #t is estimated that only 'J of the leaves in @ar*oniferous de!osits e-hi*it dama e from feedin " 9er*ivores do not make a si nificant im!act on !lant life until the Permian $Cenrick 2 =avis, %&&', !!" 1//;1/5)" .alls are e-cessive rowths on stems, leaves, cones, and flowers caused *y insect feedin or e ;layin " The earliest fossil alls are found on the !etioles of Psaronius tree ferns of the Late @ar*oniferous" #nsect all fossil diversity and a*undance takes off with the advent of flowerin !lant evolution in the @retaceous $.rimaldi 2 +n el, %&&(, !" (B)" #nsects !roduce tunnels in wood known as *orin s or alleries" Some insects eat the cam*ial layer while others eat fun us that rows within the alleries1 still others eat the wood itself" The oldest *orin s and alleries in wood, attri*uted to mites, are known from the @ar*oniferous" The first definitive *eetle *orin s are from the Triassic" There are some *orin s in !ermineralized Triassic;a ed wood from 0rizona that are attri*uted to %'

termites or *ees1 however, they may *e *eetle *orin s $.rimaldi 2 +n el, %&&(, !" (' 2 (()"

Permineralized Termite @o!rolites in Palmoxylon

Leaf mines are meanderin tunnels !roduced *y the feedin larvae of some *eetle, fly, and sawfly s!ecies" The first definitive leaf mines first a!!ear in the leaves of Triassic conifers and !teridos!erms" #nterestin ly, the a*undance and diversity of fossil leaf mines coincides with the radiation of flowerin !lants $0n ios!erms) durin the @retaceous" Leaf mines have *een used to esta*lish the !ersistence of insect and !lant associations" For e-am!le, the larvae of certain moth families have *een eatin the leaves of Quercus $oak) and Populus $!o!lars) for %& million years and his!ine *eetles have *een eatin the leaves of Heliconia for 5& million years $.rimaldi 2 +n el, %&&(, !" (%)"

@ar*onized Po!lar Leaf +-hi*itin Mar inal Feedin

@addisfly larvae live in lakes, !onds, and rivers" Many *uild distinctive !rotective cases from *its of sand, shells and ve etation" Fossil caddisfly cases can often *e identified to the family or even enus level" The oldest larval caddisfly cases $Tricho!tera) are found in the Durassic $.rimaldi 2 +n el, %&&(, !" (1)"

%(

Celliforma is a fossil *ee nest $in the form of su*terranean e-cavations) that is first found in Late @retaceous de!osits" Celliforma is found from the @retaceous to the Pliocene $.rimaldi 2 +n el, %&&(, !" (1)" Termite *orin s a!!ear in the @retaceous and re!resent the oldest undis!uted fossil nest for social insects $.rimaldi 2 +n el, %&&(, !" (')" Coprinisphaera is the fossil *urrow of a scara*aerine dun *eetle, which makes its first a!!erance durin the Paleocene" Coprinisphaera lived from the Paleocene to the Pleistocene and had a wide eo ra!hic ran e *ein found in South 0merica, 0ntarctica, 0frica and 0sia" Coprinisphaera coincide with the evolution of the first ecosystems to have a*undant mammalian her*ivores" +vidence for the first scara* tunnels are found in the co!rolites of her*ivorous dinosaurs from the Late @retaceous of Montana $.rimaldi 2 +n el, %&&(, !" (&)" Trace fossils can also hel! to esta*lish evolutionary trends" =ee! *urrows in marine sediments first a!!ear in the fossil record durin the late Precam*rian and indicate the !resence of soft;*odied coelomates $Prothero, 1334, !" %%5)" The earliest evidence for her*ivory in insects a!!ears in the @ar*oniferous" S!ecimens of %europteris and "lossopteris seed fern leaves have *een found that show si ns of mar inal and surface feedin " =efinitive leaf mines, which are !roduced only *y insects with com!lete metamor!hosis, first a!!ear in the Triassic" The first undis!uted *ee nests and termite *orin s a!!ear in @retaceous de!osits addin evidence to *ody fossils that social insects had evolved *y this time $.rimaldi 2 +n el, %&&(, !! (&;(()" Trace fossils re!resentin marine environments are used to determine animal *ehavior as well as esta*lishin the ty!e of sedimentary environment in which the rock formed" #n fact, marine trace fossils are often classified into *ehavioral cate ories1 does the trace fossil re!resent restin , dwellin , crawlin , razin or some other ty!e of feedin *ehavior $see Prothero, 1334, ! '&/ for more details): Perha!s the most !ractical way to classify marine trace fossils is *y their associations with a !articular sedimentary environment or ichnofacies" #n marine environments different ichnofacies are associated with different water de!ths, !hysical ener ies $wave 2 current conditions), or even ty!e of su*strate" #chnofacies have *ecome a standard tool for sedimentary eolo ists as well as !aleontolo ists $Prothero, 1334, !" '&/)"

&phiomorpha $Shrim!;Like 7r anism) 8urrow

#chnofossils are im!ortant tools that hel! eolo ists inter!ret sedimentary environments, !aleo*athymetry, and !rovide clues to the dia enetic history of some sedimentary rocks" For the !aleontolo ist and fossil collector ichnofossils re!resent fossilized *ehavior and !rovide im!ortant clues to !aleoecolo y and !aleoenvironments" %/

Pse"do%ossils

Man anese =endrites on Sandstone

Pseudofossils are o*6ects that do not have a *iolo ic ori in, *ut may *e mistaken as a fossil" Mineral and rock !atterns of inor anic ori in formed !urely *y natural eolo ical !rocesses may *e mistaken for fossils" =endrites de!osited *y mineral rich water !ercolatin throu h rock layers may have the a!!earance of a well !reserved !lant" @oncretions may look like e s or other o*6ects that have a *iolo ic ori in $#vanov, 9rdlickova, 2 .re orova, %&&1, !" 1&)" Concl"sion Fossils reveal that different or anisms have lived at different times" =ifferent modes of !reservation are a testament to the fact that environments have chan ed throu h time" The different life forms re!resented *y the fossil record a!!ear in an irreversi*le and thus, knowa*le order" The history of life on +arth is a true revelation from the fossil record that throws li ht on e-tinction, evolution and the continued creation of life" &ibliography 8lankenshi!, Sadekar, 2 <aymond" $%&&5)" The +volutionary Transition from 0no-y enic to 7-y enic Photosynthesis" #n Falkowski, P"." Cnoll, 0"9" O+dsP E'olution of Primary Pro ucers in the !ea" $!!" %1;B()" @hinaG +lsevier 0cademic Press" 8orror, ="D" $1344)" (ictionary of )or *oots an Com+ining ,orms" @aliforniaG Mayfield Pu*lishin @om!any" 8oucot, 0"D" $133&)" E'olutionary Paleo+iology of -eha'ior an Coe'olution" ?ew ForkG +lsevier @leal @"D" 2 Thomas, 8"0" $%&&3)" Intro uction to Plant ,ossils" Hnited Cin domG @am*rid e Hniversity Press" @ownen, <" $%&&()" History of Life O'th +ditionP" Malden, MainG 8lackwell Pu*lishin " +sch*er er, 8" $%&&&)" Coprolites" Suite 1&1"com

%5

+tter, E"" $%&&%)" 8eecherKs Trilo*ite 8edG 7rdovician Pyritization for the 7ther 9alf of the Trilo*ite" #n 8ott6er, ="D", +tter, E", 9adadorn, D"E", 2 Tan , @"M" O+ds"P Exceptional ,ossil Preser'ation. / 0nique 1ie# on the E'olution of Marine Life $1B1; 1'1)" ?ew ForkG @olum*ia Hniversity Press" .arcia, F"0" 2 Miller, ="S" $1334)" (isco'ering ,ossils. Ho# to ,in an I entify *emains of the Prehistoric Past" PennsylvaniaG Stack!ole 8ooks" .rande, L" $134')" Paleontology of the "reen *i'er ,ormation2 #ith a *e'ie# of the ,ish ,auna O%nd editionP" The .eolo ical Survey of Eyomin , 8ulliten /B" .rimaldi, =" 2 +n el, M"S", $%&&()" E'olution of the Insects" ?ew ForkG @am*rid e Hniversity Press" #vanov, M", 9rdlickova, S" 2 .re orova, <" $%&&1)" The Complete Encyclope ia of ,ossils. / Comprehensi'e "ui e to ,ossils from /roun the )orl " ?etherlandsG <e*o Pu*lishers" Danssen, <"+" $13533. Lea'es an !tems from ,ossil ,orests. / Han +oo4 of the Paleo+otanical Collections in the Illinois !tate Museum" S!rin field, #llinoisG #llinois State Museum" Dohnson, C"<" 2 Stucky <"C" $133()" Prehistoric 5ourney. / History of Life on Earth" 8oulder, @oloradoG <o*erts <inehart Pu*lishers" Cenrick P" and =avis, P" $%&&')" ,ossil Plants" Smithsonian 8ooksG Eashin ton" Martinez;=elclos, Q", 2 Darzem*owski, +" $%&&&)" Fossil insects in rocks" Meganeura )e+site" htt!GMMwww"u*"eduMd!e!Mme aneuraM(%inrocks"htm Cnoll, Summons, Eald*auer, and Lum*er e" $%&&5)" The .eolo ical Succession of Primary Producers in the 7ceans" #n Falkowski, P"." Cnoll, 0"9" O+dsP E'olution of Primary Pro ucers in the !ea" $!!" 1BB;1/B)" @hinaG +lsevier 0cademic Press" Lockley, M" 2 Meyer, @" $%&&&)" (inosaur Trac4s an &ther ,ossil ,ootprints of Europe" ?ew ForkG @olum*ia Hniversity Press" Meyer, 9"E", $%&&B)" The ,ossils of ,lorissant" Eashin tonG Smithsonian 8ooks" Miller, T" $%&&3)" #ce 8a*yG Secrets of a Frozen Mammoth" %ational "eographic, May %&&3, vol" %1(, ?o" (, !!" B';'3" Mustoe, ."+" $%&&B)" Microsco!y of Silicified Eood" Microscopy To ay, vol 11, no /, !!" B';B5"

%4

Mustoe, ."+" $%&&4)" Mineralo y and eochemistry of late +ocene silicified wood from Florissant Fossil 8eds ?ational Monument, @olorado, in Meyer, 9"E", and Smith, ="M", O+ds"P, Paleontology of the 0pper Eocene ,lorissant ,ormation2 Colora o $!!" 1%5;1'&)" .eolo ical Society of 0merica S!ecial Pa!er 'B(" ?udds, D"<" 2 Selden P"0" $%&&4)" ,ossil Ecosystems of %orth /merica. / "ui e to the !ites an Their Extraor inary -iotas" @hica oG Hniversity of @hica o Press" Pfeiffer, ="0" $%&&/), Eating ,ossil ,uels. &il2 ,oo an the Coming Crises in /griculture" @anadaG ?ew Society Pu*lishers" Piccini S" $1335)" ,ossils of the "reen *i'er ,ormation" #talyG .+7F#? s"r"l" Prothero, ="<" $%&&')" -ringing ,ossils to Life. /n Intro uction to Paleo+iology O%nd editionP" ?ew ForkG Mc.raw;9ill" Prothero, ="<" $1334)" -ringing ,ossils to Life. /n Intro uction to Paleo+iology" ?ew ForkG Mc.raw;9ill" <aven, P"9", +vert, <"F", 2 @urtis, 9" $1341)" -iology of Plants OBrd +dP" ?ew ForkG Eorth Pu*lishers, #nc" <ich P"V", <ich T" 9", Fenton, M"0", 2 Fenton, @"L" $133/)" The ,ossil -oo4. / *ecor of Prehistoric Life" Mineola, ?FG =over Pu*lications, #nc" <oss, 0" $%&1&)" /m+er. The %atural Time Capsule" ?ew ForkG Firefly 8ooks" <othwell, ."E" $%&&%)" @oal 8allsG <emarka*le +vidence of Palaeo-oic Plants and the @ommunities in Ehich They .rew" " #n =ern*ach, H" 2 Tidwell, E"=" !ecrets of Petrifie Plants. ,ascination from Millions of 6ears $!!" B3;'5)" .ermanyG =,7<7 Pu*lishers" Schluter T", Cohrin , <", 2 .re or, 9;D" $%&&B)" =ra onflies !reserved in trans!arent y!sum crystals from the Messinian $H!!er Miocene) of 0l*a, northern #taly" /cta 7oologica Craco'iensia, '/ $Su!!lement;Fossil #nsects), !!" B5B;B53" Scho!f, D"M" $135()" Modes of Fossil Preservation" *e'ie# of Palaeo+otany an Palynology, vol %&G !!" %5;(B" Seilacher, 0", Marshall, @", Skinner, 9"@"E", Tsuihi6i, T" $%&&1)" 0 fresh look at sideritic "co!rolites"" Paleo+iology, vol %5 ?o" 1G 5;1B" Selden P" 2 ?udds, D" $%&&')" E'olution of ,ossil Ecosystems" @hica oG The Hniversity of @hica o Press"

%3

Stein, @"L" $134%)" Silica <ecrystallization in Petrified Eood" 5ournal of !e imentary Petrology, vol (%, no '" !!" 1%55;1%4%" Tan , @"M" $%&&%)" 7rsten =e!osits from SwedenG Miniature Late @am*rian 0rthro!ods" #n 8ott6er, ="D", +tter, E", 9adadorn, D"E", 2 Tan , @"M" O+ds"P Exceptional ,ossil Preser'ation. / 0nique 1ie# on the E'olution of Marine Life $115;1B&)" ?ew ForkG @olum*ia Hniversity Press" Taylor, T"?", Taylor +"L" 2 Crin s, M" $%&&3)" Paleo+otany. The -iology an E'olution of ,ossil Plants O%nd +dP" ?ew ForkG 0cademic Press" Thom!son, #" $134%)" %ational /u u+on !ociety ,iel "ui e to ,ossils" ?ew ForkG 0lfred 0" Cno!f" Tidwell, E"=" $1334)" Common ,ossil Plants of )estern %orth /merica" O%nd +dP" Eashin tonG Smithsonian #nstitution Press" H"S" =e!artment of +ner yG htt!GMMwww"ener y" ovMener ysourcesMfossilfuels"htm Vinther D", 8ri s, ="+"., Prum, <"7", 2 Saranathan, ." $%&&4)" The @olour of Fossil Feathers" -iology Letters, vol 'G (%%;(%(" Ealker @" 2 Eard =" $%&&%)" !mithsonian Han +oo4s. ,ossils" ?ew ForkG =orlin Cindersley Eeitschat, E" 2 Eichard, E" $%&&%)" /tlas of Plants an /nimals in -altic /m+er" MunchenG Verla =r" Friedrich Pfeil" Eilhelm Danzen, D"$%&&%)" /rthropo s in -altic /m+er" .ermanyG 0m!y- Verla" Eilliams, =" 8", $%&&4) #ts a =irty Do*, 8ut SomeoneKs .otta =o #tG Fossilized feces reveal si nificant details a*out ancient life" Earth, Se!t"

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