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Online Publication Date: 01 September 2007 To cite this Article: Retallack, Gregory J. (2007) 'Growth, decay and burial compaction of Dickinsonia, an iconic Ediacaran fossil', Alcheringa: An Australasian Journal of Palaeontology, 31:3, 215 - 240 To link to this article: DOI: 10.1080/03115510701484705 URL: http://dx.doi.org/10.1080/03115510701484705

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The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 O:10.1080/03115510701484705 DOI: (online) Ó 1752-0754 (print)/ISSN 0311-5518 ISSN hspprpeet e esrmnsof measurements new presents paper This u only but aiiswith larities eemr ietoeo lns ihn n ug,ta fwrs elfihso anemones. or jellyfishes [[email protected]] worms, of Retallack than fungi, G.J. and of lichens compaction burial plants, and of decay those Growth, like animals. more colonial were and worms. xenophyophorans or plants, jellyfish fungi, logs, lichens, fossil than compaction-resistant more and e words: Key 23.3.06 revised 18.11.05, received USA; 97403-1272, osl Gasnr18) ait fother of as variety such fossils A 1984). (Glaessner Ediacaran fossils elongate and circular seg- irregularly unskeletonized, other disposed unlike 1A), (Fig. symmetrically ments of regularity rtl eomto niaieo yii,sdrtco actc‘et ak’o eohohrnaguiaeskeletons. agglutinate well-preserved xenophyophoran or overlapping masks’ no ‘death calcitic because or sideritic and Dickinsonia pyritic, interaction, bent of and indicative competitive Folded deformation found. brittle of been indicative ever have rims specimens reaction show specimens today, than effective less xenophyophorans. Australian decay in South of as aerobic length agglutinate-mineralization or of width or of regime in independent today, thickness impressions than Neoproterozoic in variation unskeletonized surface a great the as the to However, nearer preservation attributed much Its pyritization been mushroom. microbial has or lichen sandstones protist, quartz xenophyophoran polyp, jellyfish, Dickinsonia 31 Alcheringa R RETALLACK J. GREGORY of compaction burial and decay Growth, in fisae ecig14mi length in m 1.4 reaching 1996). age, (Jenkins the its was of It 2000). giant Shu and & (Niu 2004), China perhaps Grazhdankin (Jenkins 1992, Australia (Fedonkin al Neoproterozoic South et from of known sandstones best fossil, DICKINSONIA aebe nepee ssietal rtml rcs u a lob nepee sriiosbsso decayed of bases rhizinous as interpreted be also rings. can fairy but in tracks, of arranged wilting tumble specimens decay, mushrooms or Decayed during or trails distortion jellyfish. and lichens slime and clotting worms as crustose unlike but fossil interpreted mushroom, and been or modern lichen have of leaf, fossil shrinkage a osmotic of or wilting the like decay of nioi daaa fossil Ediacaran iconic an ETALLACK 07Ascaino utaainPalaeontologists Australasian of Association 2007 93 ekn 92,adRussia and 1992), Jenkins 1983, . .. etme,20.Got,dcyadbra opcinof compaction burial and decay Growth, 2007. September, G.J., , saNortrzi,Eicrnfsi,vrosycniee oyhee ublaino nei worm, annelid or turbellarian polychaete, a considered variously fossil, Ediacaran Neoproterozoic, a is a eitn ocmato yoebre,lk osllcessc as such lichens fossil like overburden, by compaction to resistant was Dickinsonia Dickinsonia 1-4.IS 0311-5518. ISSN 215-240. , Dickinsonia Andiva Dickinsonia sa cncEdiacaran iconic an is daaa,Peabin ahnm,growth. taphonomy, Precambrian, Ediacaran, , and scniee here. considered is Vendia Fdni 2002), (Fedonkin a striking a has , eateto elgclSine,Uiest fOeo.Egn,OR Eugene, Oregon. of University Sciences, Geological of Department hwsimi- show Dickinsonia . Dickinsonia oprsnwt h iyprstcpoly- parasitic worm tiny chaete the 1956). with Moore Comparison no & in (Harrington distin- found jellyfish other symmetry Dipleurozoa and bilateral order by 1947), guished an (Sprigg to jellyfish assigned a considered 92 eln 91 ekn 96,despite 1996), Jenkins 1991, Gehling (Runnegar sway 1982, greatest Fedonkin the had 1968, have Termier 1981) & flat- (Termier turbellarian worm or worm 1979) annelid Morris extinct (Conway an as interpretations first At problematic. main o eete preserved? they decay? were they How Did they? were organism of kind What fossils. enigmatic these about questions fundamental three of tests of as Australia South Gorge, depth Brachina and Hills Ediacara and the from length width, eelfimatcmn n iie eiiiy u no but flexibility, limited and attachment firm reveal ilgclante of affinities Biological Dickinsonia sitrrtdt essiebcueadjacent because sessile be to interpreted is Dickinsonia lososidtriaegot like growth indeterminate shows also Spinther Spongiophyton Dickinsonia Dickinsonia nioi daaa fossil. Ediacaran iconic an , Wd 92,and 1972), (Wade Dickinsonia Dickinsonia Dickinsonia and rae narcs in arrayed Dickinsonia Thucomyces Dickinsonia sevidence is was re- , , Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 B 1 RGR .RETALLACK J. GREGORY 216 i.1 Fig. ere fdcy( decay of degrees ( ( interaction Hills competitive Ediacara the likely in Quartzite Rawnsley the of ¼ 170 D F13760, . iknoi costata Dickinsonia ¼ 197 E F13977, E .Saebr r l m pcmn r oae nteSuhAsrla uem(A Museum Australian South the in located are specimens cm; 1 all are bars Scale ). ( A ¼ upper, 139 n odnMsu,Uiest fOeo (C Oregon of University Museum, Condon and F14359) B ,udfre vryn ipemr ( mark ripple overlying undeformed ), B–E and ) .tenuis D. A–B , D–E ( A oe ny rmtelt rcmra daaaMember Ediacara Precambrian late the from only) lower n rciaGre( Gorge Brachina and ) C ,wikigadflig( folding and wrinkling ), C ,soigmria aos( haloes marginal showing ), ¼ F34285). ALCHERINGA D ,addifferent and ), ¼ F17462, A ), Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 o pcfial ugl(Peterson fungal & specifically for Grazhdankin Seilacher 2001, 2002, Seilacher Reitner microbial & the generally (Steiner for of evidence parts gathering older in (Brodo thallus grazed or often are necrosed which example, lichens, is crustose for ridged but with compared, 1995), regular 1994, strikingly actino- (Retallack or fungus bacterium lichenized or mushroom symmetry. radial cnidarian Dickinsonia rather usual symmetrical the mesenteric than bilaterally its is anemone, structure or polyp parable in skeletoniza- coralline tion calcareous of trace no rlce (Yuan lichen or scleractinian The giant 1972). such coral no (Tendal in is found protists mineralized- there skeleton and the kind, agglutinate of this trace of physicochemical cell for large single unusually is a m Seilacher 1.4 although 1993, 2005), (Zhuravlev sea the deep of protists xenophyophoran with larities 1989). (Seilacher esrmnso ievrainin variation size of My measurements 2003). (Fedonkin Seilacher Proarticulata phylum or 1992, Vendozoa al et phyla (Seilacher extinct Vendobionta the in placed fossils. Ediacaran similar- superficial to ity has also mushrooms, to Dickinsonia’s a eseetv hni stdy My today. is the test it depth than impression of effective measurements decay history. less were Neoproterozoic Earth was fossils because Ediacaran in preserved that times proposed other soft-bodied He preserve at not fossils do sand- which quartz stones, non-carbonaceous, fos- red, unskeletonized in of sils solution preservation a as the (1989) for Seilacher by raised was growth. of modes ating nia ALCHERINGA h su fdcyi daaa fossils Ediacaran in decay of issue The etteevroshptee yevalu- by hypotheses various these test 03 retnttriploblastic-metazoan extinct or 2003) . Fungia Dickinsonia Dickinsonia a lobe nepee sa as interpreted been also has onmdfrisresemblance its for so-named , tal et ateslk construction mattress-like Dickinsonia Vlnie19) hr is There 1992). (Valentine Dickinsonia tal et n fi eeacom- a were it if and , 01.Teehsbeen has There 2001). . 05 ffiiisof affinities 2005) . tal et lohssimi- has also lohsbeen has also 03,and 2003), . tal et Dickinso- 2003) . tal et . EA OF DECAY iwi that is another Yet view cm) 1999). 2 – (Gehling (1 mats buried microbial shallowly mineralized within masks’ pyrite by ‘death in is as possibility leaves Another preservation of 1977). encrustation (Spicer comparable ponds oxide burial, iron after with shortly cementation or postulated before example, who for (1968), see, been Wade anomalous: long as has fossils perceived Ediacaran of paction of of which clarity impressions. clarity and depth retardation, consistent and reveal decay should depth re- vs be varying impressions, should by which vealed decay, of hypothesis esrmnso eitnet uilof burial to resistance Dickinsonia with of comparison for measurements lichens, fossil resis- of with compaction tance together the of here, measurements presented new are wood resistance of compaction the of measurements (Fletcher diagnostic (Jahren al lichens et were they that com- indicates (MacRae also isotopic of carbon quartz of position range meta- and of The veins 1999). by chlorite cut is morphic it where slabs, and Thucomyces considered myces (1976) Cloud Although Hallbauer 1974, Thucomyces totaxites Jahren giophyton Devonian including 2002), & newly (Jurina Krassilov lichens of and fungi compaction fossil burial recognized with time first kltn r itecmatdb burial but by 1994), now (Retallack compacted wood with little compared are (Retallack mineral skeletons or lichens car- Cements 2002). biopolymer (Fedonkin of coherent apace a chitin perhaps or 1994), lignin wood with of comparable biopolymer, tough h rsre eitnet uilcom- burial to resistance preserved The 03,ee faslt ausaeun- are values absolute if even 2003), . Dickinsonia ob natfc fai maceration, acid of artefact an be to tal et Hee 01,adArchaean and 2001), (Hueber (Stein . 03,Siluro-Devonian 2003), . Dickinsonia skonwti thin-sections within known is Hlbur&VnWarmelo Van & (Hallbauer DICKINSONIA pnipyo minutissimum Spongiophyton tal et tal et a ecmae o the for compared be can tal et 97 aRe1999). MacRae 1977, . 19,Rtlak1994, Retallack .1993, 04.Additional 2004). . a oekn of kind some had Thuco- Spon- Pro- 217 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 . m(i aaeMuti) . km 3.6 Mountain), (Cabin km 3.3 Savage and Mountain), Creek (Sugar (Big km 7.8 Beaumont of thermo-mechan- modelling logs from ical estimated fossil been America have North studied from 1995) 1994, previously (Retallack for of Depths burial 1950). (Taylor grain per neighbour- grains of ing supported number esti- the 3), counting geological point (Table by overburden from of derived mates was of Depth burial fossils. Ediacaran like quartz sandstones in preserved Chaloner commonly & are (Rex and 1983) sandstones width compressed not are but logs thickness quartz in because 3), in – 2 (Tables compressions fossil of 1995) width log and 1994, thickness the (Retallack measuring studies phonomic hssuyi ae neaiaino type of examination of on collections based is study This methods and Materials hndtiso ibn in ribbing of details than and uncommon) 1966 Wade, common), most the costata species: four Dickinsonia recognized who (1992) of Jenkins classification the follows Condon Flinders study This (Retallack the 1994). Oregon Gorge, in of University Australia, Collection, Brachina South speci- Ranges, from two and and Adelaide, mens Museum of of University Geophysics Australian the and Geology South of Department the Australia, in South Ranges, Flinders and Hills 1 RGR .RETALLACK J. GREGORY 218 mi imtrfo lnrsurface planar a from diameter 10 piston in a of mm 3 protrusion a measures of It flatness of surface. high-precision engi- degree the automotive determine a to neers for A is designed 1). gauge instrument (Table depth mm im- 0.025 the Vernier to of accurate depth depth Vernier gauge a the with and measured was mm, pression 0.1 accu- to micrometer Vernier rate were a specimens all with of measured length and width The 6 hsppretnspeiu isota- previous extends paper This m opcsotgnrllvl rather levels general out picks so cm, 1 .rex D. , Dickinsonia .lissa D. .tenuis D. pig 97(yfar (by 1947 Sprigg, ekn,19 (both 1992 Jenkins, ae 92(rare). 1972 Wade, Dickinsonia rmteEdiacara the from tal et lese & Glaessner 18)as (1988) . . 2001). ncosscinwti permineralized within (Hallbauer cross-section specimens in al et h atnn ftblrhpa nSEM in Hallbauer hyphae (Chaloner from tubular 4) photomicrographs of fungi (Table flattening estimated fossil the was of lichens compaction and Burial Jahren 2001, 2003). Hueber 1999, (MacRae decade last the within confirmed or discovered Dickinsonia Ediacaran Gehling for by advocated (1999). burial m) 1500 – (400 of fossils include depths thus data shallow new These to and 2). sand- (Fig. depth quartz stones chosen burial within of variation were range petrographic here wider a studied explore logs the fossil 1995), and 1994, (Retallack of matrix fossils burial Ediacaran likely for maximal to close chosen depth burial logs fossil studied previously Unlike Corner). Bragg and Creek o edndb ae olfrainfrthe for formation soil South later and by of brown, reddened Neopro- or not red shales originally Red were Cambrian Australia and sandstones. and terozoic white shales red flaggy mud-cracked, and marked nia growth-series of this analysis. assumption justify the further necessary and paragraphs lived, they following where preservation bed- evidence for is itself same in This series the 1A). (Fig. on planes ding size in- growth different with of 1994), dividuals complete Retallack 1982, be (Runnegar remarkably to appear 1996). a collections Jenkins Australian 1972, South (Wade long mm 1400 Dickinsonia measurements length and width by indicated Affinities locmae ihpeevto of preservation with compared Also h rsrainlhbtt of habitats preservational The a eifre rmecoigripple- enclosing from inferred be can 91,bcuetetbsaecircular are tubes the because 1991), . tal et r oslfniadlichens and fungi fossil are pcmn ag rm4to 4 from range specimens 97 tohr18,Gensel 1988, Strother 1977, . tal et ALCHERINGA 97 Hueber 1977, . tal et Dickinso- .1974, tal et . Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 a aiswt bnatelephant-skin- abundant mats, microbial with of texture facies tidal in Even flat observations. environ- following the subtidal on based autochthonously to preserved and estuarine ments of ganism sandstones flaggy the of 1994). soles (Retallack the on mon ee ecinrmt h nrahetof encroachment the thic- to a rim has reaction specimen kened right-hand The shows of mens. 1B) pair known (Fig. closest the here illustrated (Gehling specimen pre- fossil clearly preserved another overlap served to known storms Hagadorn is by 1994, al mudflats Retallack et and 1989, beaches thrown (Norris range on size up uniform very of of schools stranded jellyfish unlike size, in vary mens 1996). in sub- (Jenkins shorefaces deposited shallow tidal and flaggy probably palaeochannels the estuarine were and floodplains, in sandstones down and laid flats likely most tidal were that 1999). beds (Gehling red like The evaporites cracking sulfate microbial in and found pa- from deformation exposed mat come periodically of laeoenvironments evidence and Further oxidizing 1996). inactive slow (Jenkins tectonically coast and a along water quartz, accumulation well-oxidized of entirely indicating of almost well- clean grains, and well-sorted, rounded unusually with and matter, but organic white clasts, mostly red have are also sandstones flaggy etwt te niain flt Precam- late (Schmidt of palaeopoles indications brian other consis- with are in tent and tests deformation fold soft-sediment palaeomagnetic pass Hematite 1990). cements (Moore from minerals radiating iron-rich stain diffuse a red. is are pigment breccia Red intraformational Clasts in 1967). shale Curnow of the & below m (Goldring 91 surface of depths to boreholes is in Member red Ediacara The reasons. following ALCHERINGA Dickinsonia 02.N well-preserved No 2002). . Dickinsonia Dickinsonia a rbbyassieor- sessile a probably was rayohrclearly other any or , mrsin r com- are impressions tal et Dickinsonia Dickinsonia tal et tal et 93 Jenkins 1983, . 93.The 1993). . Dickinsonia 05.A 2005). . speci- speci- EA OF DECAY ylnal19) n rsoelichens crustose and & (Brodo (Kidwell 1998), bryozoans Gyllenhaal Buss encrusting & (Jackson 1975), sponges and allelo- ad- corals between to jacent interaction similar competitive specimen, pathic left-hand the et aesonthat shown experi- Flume have stac- corpses. for ments case unattached the of be against king would shuffled as another, nor one buckled, nor contact, rl tahd n npaeo growth. of place in and attached, but firmly deposition, during Dickinsonia tran- sorted been and have sported that (Gehling fossils for fossils appropriate of al transport et other or of sorting marked specimen size evidence marks, a tool show of Museum absence slabs mm Ediacaran 9 – Australian P41164). 2 by halves (South undeformed two rated rc hog one mud- A through attached. crack apparently sediment, of al of et Dismem- cross- specimens 1998). of Baumiller bered base & the (Schopf at beds preserved be in to substrate the order to attached firmly been have irb n vlsae n (2) and shape, oval and midrib, costata (1) species: inter- nia distinct here two origin, as the preted though arrays linear 1). (Table here available presented results the becoming as are (Droser planes single from Collec- bedding populations samples. individual of of tions suite the are available but largest population, unlikely single a are a Hills represent from Ediacara to the largely in years site single many over collected addeapeo h aeseisof species same ex- the two and of these contracted example a of panded inter- than co-occurrence here rather a illu- species, is as widely 1A) A preted (Fig. 3A). and specimen (Fig. midrib, strated wide shape segments, elongate narrow with h pcmn of specimens The rwhi it n eghso two show length and width in Growth 05 g )so xesv disruption extensive show 9) fig. 2005, . 05.Aayi fgot eisi not is series growth of Analysis 2005). . tal et tal et ihwd emns narrow segments, wide with , 03,btntyta numerous as yet not but 2003), . per ohv ensessile, been have to appears 01.Term r o in not are rims The 2001). . DICKINSONIA Dickinsonia Dickinsonia Dickinsonia Dickinsonia a sepa- has .tenuis D. Dickinso- (Gehling casually must 219 , Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 1308. 150.737 0.991 4.140 71.5 2.210 0.356 16.5 1.346 2.235 1.600 193.7 2.261 56.7 1.676 83.7 19.3 1.676 18.7 22.8 0.152 49.8 106.7 1.753 2.515 323.8 Incomplete 140.7 59.7 0.305 17.9 53.8 Incomplete P14350 0.965 22.5 17.6 P14344 1.626 61.9 44.5 P14342 0.584 208.6 P14334 0.737 89.6 41.3 P14333 2.261 53.3 36.9 P14331 2.311 21.5 21.7 P14330 43.5 28.1 P14328 2.311 61.3 165.9 P14327 2.007 48.7 63.5 0.330 P14322 1.626 37.3 72.6 P14241 1.600 30.7 P14221 1.372 38.5 25.1 0.9 P13799 63.3 45.5 P13767 41.7 72.3 25.7 1.753 P13760 99.5 42.4 P13760 61.6 0.838 P13718 33.7 1.600 Incomplete P12900 2.057 55.8 1.422 28.5 P12749 33.7 2.591 P12729 1.905 38.5 0.279 31.3 P12728 65.9 36.7 P12727 81.4 23.9 P12725 2.642 37.5 61.8 P12724 1.549 32.6 1.194 64.7 P12690 Incomplete 0.838 50.6 0.203 P12678 Incomplete 0.940 P12557 0.864 34.2 147.9 MW741a,b Incomplete 54.5 0.660 74.3 21.7 MW563 1.626 51.6 1.2 50.8 8.2 MW561 18.8 MW559 41.5 MW558 169.7 Incomplete 1.626 34.6 3.505 MW557 25.1 35.5 MW556 Incomplete 1.702 2.210 68.2 MW554 8.7 MW553 55.1 64.7 MW299 63.7 46.2 MW205 2.600 60.6 61.2 95.9 4.000 MW204 MW199 MW196 Taxon 78.5 MW189 77.9 Incomplete 44.9 (mm) Height MW187 132.8 161 MW168 (mm) Depth MW165a,b MW1275/1 Incomplete (mm) Width F17462-4 Incomplete F17462-4 (mm) Length CCF34288 CCF34285 Number RETALLACK J. GREGORY 220 al 1 Table esrmnsof Measurements . Dickinsonia osl rmSuhAustralia. South from fossils iknoi costata Dickinsonia costata Dickinsonia tenuis Dickinsonia rex Dickinsonia rex Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia tenuis Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia tenuis Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia ALCHERINGA ( continued ) Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 nubrd3. 521590.9 specimens). 1.295 two in 1.473 only known 1.118 latter 1.549 4.674 (this 0.305 counterpart convex the two (1992). of Jenkins and is surface after height 80.3 Museum, and is the and impression Australian Taxonomy concave axis from of South 2.159 long surface of below the distance 65.3 distance dimension 1.397 is in is the depth 49.9 Length whereas F-) Oregon. axis, short (P-, of 2.896 35.2 of University 131.7 collections dimension 0.305 Collection, width 32.2 bulk Condon the and in (T-), (CCF-) 0.991 specimens Sprigg Reginald by 1.600 and 91.7 87.5 2.007 Note Incomplete 58.8 62.3 82.6 1.397 55.6 23.8 0.457 35.8 40.1 Unnumbered 45.5 2.464 Unnumbered 60.6 4.013 Unnumbered 65.8 1.295 101.9 Incomplete 68.5 Unnumbered 25.5 34.5 30.2 T61;2061 2.311 0.889 25.7 50.8 T60;2054 0.457 81.7 60.8 T54;2050 0.432 43.2 95.9 T53;2004 0.152 35.5 T51;2000 2.311 49.1 100.1 23.9 T50;2001 1.499 41.4 52.8 T47;2052 1.981 38.6 160.2 T46;2009 45.9 55.1 T45;2001 1.295 42.7 125.8 T45;1005 28.7 36.7 1.295 1.321 P21155 1.778 59.9 40.2 P18888 54.6 0.279 P17998 59.3 21.9 0.381 P14395 0.457 57.8 45.7 P14393 40.2 0.356 47.9 36.8 P14389 0.965 44.5 26.9 P14379 1.981 20.5 P14378 0.711 19.2 31.7 Incomplete P14377 0.330 24.7 62.3 P14373 43.9 22.2 P14372 4.674 0.254 19.3 34.8 P14370 0.178 33.9 29.1 P14369 2.210 34.7 22.8 P14368 0.737 26.2 P14367a 0.991 21.5 160.7 10.8 P14366 21.8 42.7 P14365b 43.9 56.7 P14365a 28.3 12.5 P14364 13.9 177.5 P14361 11.2 P14360 Taxon 59.5 P14360 65.5 (mm) Height P14359 13.3 P14355 14.3 (mm) Depth P14354 P14354 (mm) Width P14353 P14352 (mm) Length P14352 P14351 Number ALCHERINGA hsicue olcin yMr ae(W)i h elg eateto dlieUniversity, Adelaide of Department Geology the in (MW-) Wade Mary by collections includes This : al 1. Table ( Continued EA OF DECAY ). DICKINSONIA iknoi rex Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia rex Dickinsonia costata Dickinsonia tenuis Dickinsonia tenuis Dickinsonia tenuis Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia tenuis Dickinsonia tenuis Dickinsonia tenuis Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia costata Dickinsonia tenuis Dickinsonia costata Dickinsonia costata Dickinsonia 221 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 30 odnGoe South Grove, Golden L3003 30 roe igna S:creek USA: Virginia, Brooke, L3002 30 xadeCno,Utah, Canyon, Axhandle L3001 30 armta e ot Wales, South New Parramatta, L3000 ubrLclt ecito omto g References Age Formation Antarctica: Mountain, Portal L2998 description Locality number Locality RETALLACK J. GREGORY 222 .costata D. orientation. or decay satiation, to differentiating costata wider feature insonia consistent the width a Midrib is in specimen. inflated and than ribbed specimen, supposedly and contracted ribbed narrow the in 1996), wider (Jenkins is creature bodied annelid soft an other of or gut Furthermore, a larger as larger. interpreted the be midrib, as the should wide rim as its ribs one, as with specimen and the and as a large from If smaller contracted one specimen. smaller the ‘contracted’ supposed than supposedly wider rim’ a ‘contraction has supposedly growth larger specimen different the ‘inflated’ but on 3A), fall (Fig. they arrays do 1992). (Runnegar only count Not rib and age similar itgaso it n eghin length and width of Histograms S34.7811 Road: Tree One on quarry large of margin northwest Australia: N33.38687 629: and of 608 junction highways of west m 200 N39.40268 mouth: canyon at the ridge northeast the of base the above stratigraphically m 118.7 USA: E151.01289 S33.78392 School: The Kings of part northern in creek above track on Australia: E159.39183 S78.11007 spur: eastern dolerite on lower above m 34.3 r ohsrnl negatively strongly both are from 8 al 2 Table E138.73932 .tenuis D. 8 8 8 8 W77.380700 W111.68204 e oaiisfrfsi osi urzsandstone. quartz in logs fossil for localities New . ounrelated so , 8 8 8 8 8 Dick- odnGrove Golden Patapsco ot Horn North Hawkesbury elrCoal Weller beds Formation Formation Sandstone Measures rwht sxa rsxa erdciesize (Gand reproductive determinate sexual or show asexual to also growth jellyfish and Polyps very growth, from distinct determinate allometric show ooilognsswt indeterminate such worms living as free with contrast, In growth. other organisms and bryozoans colonial sea- lichens, like trees, sediment, by weeds, covered significantly until highly grow to a 4B), distribution. (Fig. skewed length modal mm the 50 times of 28 is 1996) (Jenkins largest Rtlak19,Peterson animals 1994, colonial (Retallack growth, and plants fungi, indeterminate of typical observations isometric both growth, indicate of place popula- in are tions these Because 4B). (Fig. skewed pnhrarcticus Spinther tal et Dickinsonia oeeMClu (1990) McCallum Eocene Early aecn ite(1988), Hintze Paleocene Middle i-ema Collinson mid-Permian 96.Lk otmetazoans, most Like 1996). . Cretaceous Triassic Dickinsonia and Dickinsonia t10 mlong mm 1400 at eesdiversicolor Nereis Rnea 1982). (Runnegar tal et ol Hickey & Doyle oahn(1980), Conaghan ALCHERINGA rc (1999) Bruce & Powars (1976), (1994) Talling (1980) Crawford (1980), Herbert (1994) 03.The 2003). . continued tal et tal et tal et . . . Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 igeseie of are specimen single counterparts example, a and for 1999): (Gehling parts collected of rarely best weathering slabs, by the loose enhanced Because are beds. 1.84 the specimens overlying on of concave 3.18 are soles specimens 1.06 Gehling Most 1994, 1999). 1.69 (Retallack fossils associated (on other Ediacaran 1.55 unlike impression bed), 1.18 raised underlying a the as 14.42 preserved 2.2 1968) 22.06 2.53 12.99 1.57 28.27 1.34 1.16 11.01 6.69 1.759 2.23 1.759 29.62 1.759 24.22 1.56 1.759 17.48 1.75 1.69 13.64 1.759 1.92 1.01 1.759 11.21 Dicot? 1.18 1.52 1.759 23.41 Dicot? 3.12 1.759 Dicot? Dicot? 2.03 12.77 1.5 4.11 Dicot? F36244H 23.33 1.5 Dicot? 7.52 F36244G 16.4 1.5 Dicot? 5.96 F36244F 22.32 1.5 Dicot? F36244E 18.66 L3001 F36244D Conifer? L3001 9.53 F36244C 0.5 Conifer? 1.27 L3001 F36244B 0.5 Conifer? 1.27 L3001 F36244A 0.5 Conifer? L3001 1.27 F36243D 0.5 L3001 F36243C 0.5 Glossopterid? L3001 F36243B 0.066 Glossopterid? Dicot? L3001 F36243A Dicot? Glossopterid? L3000 F36241C Dicot? L3000 F36241B Dicot? L3000 F36241A Dicot? L3000 F36247 Dicot? L2998 F36247 L2998 F36247 L2998 F36247 Taxon L3002 F36247 L3002 F36246 L3002 L3002 L3002 Specimen L3003 Locality ALCHERINGA SMP16ab n n of one and P41166a,b) (SAM like Dickinsonia measurements depth by indicated thickness of Decay size modal a soon near ceases maturity (Peterson then sexual and after slows growth their ocv ibdmr ntesl fthe of sole the a on is side mark upper ribbed the concave cases, both In P40200). Tribrachidium tal et sa‘eitn’Eicrnfossil Ediacaran ‘resistant’ a is 2003). . al 3 Table and e aao opcino osllg nqat sandstone. quartz in logs fossil of compaction on data New . Parvancorina iknoi costata Dickinsonia .rex D. (Wade (SAM et (km) depth EA OF DECAY ontso la rwhary,btinstead but arrays, growth clear show not do n t usqetcmato u to due compaction subsequent burial. its total thickness and estimating the in from from impressions the of subtracted sand depth of was by This portion supported below. a was and impression unifacial, mid- was and Thus, bing ribbing with 1999). (Gehling distinct friable, line less more and is fainter convex which its from counterpart, fine distinct concave a hematite, with of upper patina finished smoothly The is 1999). hematitic impression some cases (Gehling few a clay in grains and cemented base mm its 3 loosely at as contains 3). gap much (Fig. This as counterpart and to part My between 0 thick from 1994). range a Retallack fossils that is underlying 1968, demonstrate the measurements side of (Wade top under the bed the on mark and convex bed, overlying Burial hcns esrmnsof measurements Thickness DICKINSONIA o width Log (mm) Dickinsonia hcns (mm) thickness Dickinsonia Dickinsonia Log rib- 223 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 .5Bertrand-Sarfati 1.45 .5Bertrand-Sarfati 1.45 (km) depth Burial RETALLACK J. GREGORY 224 esls eoebra.Tefis ise to tissues first The burial. before loss thick- and ness decay leaves variable show which fossil 2001), (such lichens of or as fungi and that 1985) (Ferguson with comparable length. than and rate width in in thick- growth constrained in more growth though indeterminate ness, supports of curve idea skewed the of unimodal stages A different decay. in fossils different with of age the planes biomat from bedding polymodal expected of or contribution be Bimodal would and 4A). distributions thickness of left-skewed, (Fig. a that length shows like which curve 3, equa- unimodal growth Fig. a of from tion predicted width for each thickness the unlike bimodal, preserved is of thickness distribution size with The specimens ribs. thin weak to ribbing specimens marked thick with from vary but measured distinct, comparable specimens were the of All 3B). fossils (Fig. width for variability great 84Kn 18)04 .129 0.11 0.47 (1980) Kent 18.4 212Beaumont 12.192 212Beaumont 12.192 Poole 3.784 Nematothallus hspteno hcns aito is variation thickness of pattern This uildepth burial to Reference al 4 Table tal et tal et tal et (1991) . (1991) . esrmnso opce irsrcueo oslfniadlichens. and fungi fossil of microstructure compacted of Measurements . fSrte 98 Hueber 1988, Strother of 17)04 .930 0.09 0.48 (1976) . tal et tal et 18)04 .45 0.14 0.44 (1988) . 18)04 .54 0.05 0.48 (1988) . height/width ftb cells tube of Mean .500 11 0.07 0.45 .801 4 0.10 0.48 deviation Standard fmean of 02 rwrs(owyMorris (Conway worms or 2002) hikg,wligaddscain which desiccation, and wilting shrinkage, of thinning and Dickinsonia decay observed wrinkled The and skeins. fragments, exploded masses, and irregular torn clotted form which 1982), ihn n ug itaddcybefore decay and wilt (Brodo hyphae fungi mycobiont of (Ferguson and tissues outline hymenial lichens and last Phycobiont the 1985). as tracheids itself skeleton’ of cuticle ‘leaf a the leaving breached, is then and thins cells, mesophyll and palisade parenchymatous are leaves in decay iig(ors18,Buo 91 n fossil (Gand and 1991) jellyfish Bruton 1989, of (Norris that living from distinct is deformation, modest 3. Fig. in shown equations the growth the by and specimens, thickest represented burial- is but specimens undecayed compacted of Thickness burial. Dickinsonia of thickness in variability the perspective, this hspteno ea n hnigwith thinning and decay of pattern This esrdTaxon measured Number fcells of sas itntfo osmotic from distinct also is seiec o ea before decay for evidence is tal et 96 Hagadorn 1996, . pnipyo nanum Spongiophyton pnipyo nanum Spongiophyton Thucomyces Prototaxites Prototaxites Spongiophyton 17,p.II g 24) fig. III, pl. (1977, 17 l 2,fi.1) fig. 124, pl. (1974 Chaloner Hallbauer lichenoides 18 g 8-4) fig. (1988 18 g 8-2) fig. (1988 etfi.5D) text-fig. Gensel minutissimum 7) fig. 121, pl. (1974 Chaloner tal et ALCHERINGA 01.From 2001). . tal et tal et tal et p Strother sp: p Strother sp: : tal et (1991 . : . . . tal et tal et : : . . Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 ALCHERINGA pcmn fmrn netbae.Thick history invertebrates. natural marine dried of and twis- specimens pickled and in folds, ting wrinkles, prominent form from are data All diagrams). (pie strength mechanical thin-sections. of petrographic indications counting as point composition and grainsize and (histograms), of compaction burial 2 Fig. ergahcdt nmtie ffsi osue ocntutcmato uv flg o oprsnwith comparison for logs of curve compaction construct to used logs fossil of matrices on data Petrographic . Dickinsonia Fg ) hwn ri otcsprgana nidcto fbra compaction burial of indication an as grain per contacts grain showing 6), (Fig. EA OF DECAY tae eomirglryuo et or death upon sub- irregularly distress. the deform to strate, untethered bodies, gelatinous DICKINSONIA 225 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 2 RGR .RETALLACK J. GREGORY 226 usqetyrcgie oslfniand fungi fossil of compaction recognized burial subsequently with comparison also and by 2001), Retallack by & (Sheldon here algo- rithms compaction quantified subsequent ob- of further application previous are My servations 1994). and logs (Retallack fossil like leaves much km, 5.8 – 1.5 been buried have im- that sandstones strong quartz in remarkably pression a make undecayed, of specimens Some measurements depth by indicated resistance compaction Burial decay. of effects by isometric, obscured and is thickness indeterminate in was growth length but Growth and below. width from sand the in with of filled are portion impression the squares raised showing Open counterparts, of Australia. specimens South Ranges, Flinders tenuis D. of specimens 3 Fig. aito nwdhv eghadtikesof thickness and length vs width in Variation . coe ice)fo h daaaHlsand Hills Ediacara the from circles) (closed iknoi costata Dickinsonia Dickinsonia oe ice)and circles) (open presumably , atndt lissi opeso speci- compression (Chaloner in mens ellipses to flattened Gensel analogs modern and (Hallbauer specimens perminer- in deformation tubular alized be and from to known inferred fungi hyphae of be fossil can of lichens Compaction lichens. uvso i.3 l uvsso h toglf kwof skew left growth strong the inferred show growth. curves the indeterminate All 3. using Fig. of decay curves for corrected nesses costata Dickinsonia 4 Fig. hi it u o hcns ihburial with thickness not maintained but and width objects, their cylindrical they Spodosols that assuming were Compaction estimated is and 2001). logs fossil of Retallack Andisols & (Sheldon low- soils, as 5) curve density compaction (Fig. same depth the follows with microstructure lichen itgaso it,lnt,adtikesof thickness and length, width, of Histograms . tal et tal et 91.Tecmato of compaction The 1991). . losonaepeuilthick- preburial are shown Also . 97 ubr20) but 2001), Hueber 1977, . tal et 94 tohr1988, Strother 1974, . ALCHERINGA Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 ALCHERINGA r nw piedw rfle nhalf. in folded living or down free upside were known No are 1998). they Baumiller have & if (Schopf should them marks entrained ripple inferred overlying Currents substrate. from of the most to of body the attachment margin, the firm at suggests mm which 4 maximum only overlap, the of shows extent 1D) (Fig. here over figured folded slightly (Gehling found themselves been speci- have Many mens 1C). (Fig. sediment foresets covering 1D) of ripple-marked undeformed (Fig. (2) themselves and over folded specimens partially wrinkled ground- (1) low organisms: neither were hugging they was that suggest it vations obser- Two because shape. in spherical nor difficult, ellipsoidal more is 2001). Retallack (Histosols & coals Sheldon and of burial peats woody of as depth 5) (Fig. increasing the with follows curve logs same fossil of Frac- compaction 1983). tional Chaloner & (Rex compression and fungi fossil of deformation microstructural from palaeosols, Vertisol and in jellyfish ( dykes logs, clastic lichens lichens, of palaeosols, folding ptygmatic of from depth with compaction observed 5 Fig. hcns oprdwt sue rgnltikeso mfrteEicrnfossil Ediacaran the for cm 1 lichens. of and fungi thickness fossil original as assumed compaction-resistant with compared thickness siaigcmato of compaction Estimating rcinlcmato rdce rmsadr qain fSednadRtlak(01 oprdwith compared (2001) Retallack and Sheldon of equations standard from predicted compaction Fractional . hcmcs pnipyo,Prototaxites Spongiophyton, Thucomyces, tal et 05,adteone the and 2005), . Dickinsonia Dickinsonia ,fo opesdtikeso osllg,adfo current from and logs, fossil of thickness compressed from ), EA OF DECAY osllg Fg ) h -mcs sa upper an is case 1-cm The 5). than (Fig. logs compaction-resistant fossil more much and ato eitn sfsi ug n lichens and fungi ( fossil 4) as resistant (Fig. paction thicknesses that present shows to that of thickness comparison A cm. 1 than thicker been udn(olwn loih o quartz Retallack so & for 2001), Sheldon of Spodosols algorithm and sand over- (following of km 5.8 burden by cm compaction 1.4 before been thick that have would or bed 8-mm-thick An top, burial. during them the around swirled that through sediment evidence no protruded with mm, they 8 as thin as slabs (Retallack 1994). fungi subsur- some like to rhizomorphs attached face or rhizines or lichens, by oysters crustose rooted like like 1992), substrate (Valentine corals the to specimens cemented dismembered is partly disruption (Gehling sediment in of visible zone marked A rttxts Spongiophyton Prototaxites, Dickinsonia tal et Dickinsonia Dickinsonia Dickinsonia DICKINSONIA 05.Te eeeither were They 2005). . osl aebe on on found been have fossils opcinwsmeasured was Compaction . Dickinsonia r nieyt have to unlikely are a tlata com- as least at was , Thucomyces hc a as was which , 227 ) Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 nl ebe etr ugssaspicular a Zhu 1994, suggests (Geyer skeleton texture pebbled their not finely and were echinoderms, eldoniid they but that jellyfish indicates Ediacaran gut to U-shaped similar (Samuelson also size and fossils Morocco relief a of have sandstones Ordovician with Ernietta RETALLACK J. GREGORY 228 opcinmr ietoeo lichens, of colonial burial or those and xenophyophores, algae, like decay plants, and more growth compaction of modes ( fossils Ediacaran (Hagadorn in throws al death et jellyfish sediment during of Cambrian casts ingested natural of from came relief fossils cases, other around found been siderite. not haloes have ferruginized by or nodules sideritic mineralization Such from gained fossils strength these of some ferruginiza- that suggest and tion significant cracking local synaeresis their have but radial relief, 1994) unmeasured) (though (Cherns Sweden limit: ymaueet of measurements My of taphonomy and affinities of indications Other Rtlak19) abinjlys from nodules, (Hagadorn jellyfish Wisconsin siderite Cambrian 1994). in (Retallack compaction-resistant preserved were themselves which 5) (Fig. 5). clastic to (Fig. folded un- used dykes ptygmatically by Vertisols their equation as ravelling compaction such the soils, calibrate most as that show also Dickinsonia 2001) Retallack & (Sheldon 1994). walls (Retallack cell fungal of chitin the as such polymer, life. in thinner Presumably if resistant compaction more opce hnjellyfish than compacted el ple opcinalgorithms compaction applied Newly 02.Sc nenlmud r known are moulds internal Such 2002). . Dickinsonia Dickinsonia fGasnr 94 .8) u not but 82), p. 1984, Glaessner, of a o scompaction-resistant as not was Dickinsonia Dickinsonia upsdmdsefrom medusae Supposed . ol aebe even been have would tal et Dickinsonia tal et Protoechiuris 01,bttheir but 2001), . a togbio- strong a had seel asherae Essexella Dickinsonia tal et Dickinsonia a uhless much was 02 and 2002) . 2002). . indicate and .In 02.I h eohohr interpreta- Seilacher xenophyophore by championed the tion In 2002). (Fedonkin carapace be- dorsal from chitinous extending a foot neath 1956), flexuous soft Moore a as & or Harrington jellyfish 1947, of (Sprigg tentacles as interpreted been also the at chemicals surface. allelopathic and mats decay of advanced microbial dispersion with or persist not sediment would scale, in same marks the of and grooves leave should falsify ribs retracting because observations hypothesis, retraction These the 1E). sur- (Fig. microbial decayed obviously faces well- extensively less as on well mats specimens in as microbial 6), 1A, as (Figs clear regarded elephant-skin-tex- surfaces tured are on specimens preserved haloes 6B). (Fig. specimens Marginal lineation the small and finer than large both have thinner on but and impressions, fainter main haloes only marginal not pre- are The not are variants. species, and different servational as 3A) here (Fig. soft- regarded arrays gelatinous growth a on different fall specimens of two These as creature. expected bodied 1982), slab be Runnegar same 1972, would the Wade on 1A; and a count (Fig. segments segment from wider same with the shrunken one thinner like as with creature specimen regarded a segments a fully and to size, of desiccated smaller were impressions that animals as hydrated interpreted been pcmn htso lal preserved clearly from a comes views show three that principal all specimens The to pseudopodia. objection be could they agnlhlsaround halos Marginal increment? growth or pseudopodia tentacles, rim, Retraction and affinities the of taphonomy concerning following published arguments the of variety a perspective, jelly- reevaluate paragraphs this or polyps From worms, fish. of than organisms h lmnosmria aoshave haloes marginal filamentous The Dickinsonia Dickinsonia . ALCHERINGA tal et (2005) . have Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 ALCHERINGA pedgs etce rfe,btthe but feet, or tentacles are delicate appendages, more carapaces without preserved molluscan commonly jellyfish, and Protist, 1E). arthropod (Fig. portion decayed central extensively yet margin, filamentous enlarged in construction microtubular extensive showing Australia, South ( Gorge, portion Brachina in Quartzite Rawnsley 6 Fig. necpinlylreseie of specimen large exceptionally An . B .Cno uem nvriyo rgn(F34285). Oregon of University Museum, Condon ). iknoi costata Dickinsonia EA OF DECAY nrmn falce Rtlak19) This 1994). (Retallack growth lichen future a of a increment for foundation rhizinous a rsraino etwtotcrpcsis carapaces without unknown. feet of preservation ( A rmtelt rcmra daaamme fthe of member Ediacara Precambrian late the from ) e nte nepeaino h aois halo the of interpretation another Yet DICKINSONIA 229 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 hikg rpsdfraseie illu- specimen double Gehling a by to for strated explanation proposed alternative shrinkage an which an is specimens, been damaged heavily enabled have of but repair only defense, not advanced would allelopathic 6B). rim (Fig. like fossil a the halo of Such part showing marginal main the the specimens of those in other features an and contact tubular to actual 1B), of rim (Fig. short reaction specimen a a adjacent by showing suggested specimen especially is interpretation scmail ihte‘nu architecture ‘pneu’ the and with 1973), compatible (Pflug is Namibia of Ediacarans Peterson al 2002, et Narbonne & Newfound- (Clapham Point, ‘spindle’ land of Mistaken impressions at in Ediacarans seen is tubes branch- ing linked laterally from of construction Compar- able fossils shale. in Ediacaran preserved Newfoundland rangeomorph in nia impressed little presumably deeply and 3B) Fig. wide), decayed. see is mm: cm (4 6) (16 (Fig. features large these lichen speci- showing The like best splay rhizomorphs. men fossil, fungal and the or the rhizines from branch in away but tubes radially ribs, halo, broad fabric marginal the felt-like to a The orthogonal create diameter). mm structures 1 – tubular (0.5 are large they too because much hyphae 6). actinobacteria, individual or (Fig. be fungi of to ribbing unlikely are conspicuous These and grains than sand different than finer several coarser all of grades size structures tubular branch- ing abundant show specimens large but Australian South of Dickinsonia matrix sandstone The construction Fractal-tubular decay (Brodo and centres death their after of common is margins lichens from of regeneration and Persistence 3 RGR .RETALLACK J. GREGORY 230 hs uua etrso large of features tubular These r ietoefudb abne(2004) Narbonne by found those like are 03,i emnrlzdpetalonaman permineralized in 2003), . rcue itlgclstudy, histological precludes tal et tal et 20,fi.8). fig. (2005, . 2001). . Dickinso- akshrclcls( 5 – (4 cells mm spherical 5 – 2 (1 – dark (0.1 cells tubes woven, tubular densely hollow of constructed are the diameter) also (Retallack that permineralizations replacement show Namibian other later 1994). than or rather fungal histology, original pre- serve Namibia that of confirm Ediacarans thus permineralized (2004) discov- Narbonne New by (1989). eries Seilacher by proposed nlcea r nerdfor inferred are intest- caecae tubular inal large by Very outlined (2004). as Narbonne a construction of tubular portions unravelled fractal be the could tubes segments. The by the across run they anyway because polychaete worm, or annelid unlikely an of model be biological caecae. would intestinal as This them interpreted who decayed a in Dickinsonia illustrated been have diameter flces(ealc 1994). (Retallack podetia cells) lichens spherical of (tubular like (dark phycobionts mycobionts and most cells) and tubes), any of plant, (megascopic that or unlike is animal histology This 1994). uua osrcinmdl hscould others which this from axis central model, the Pflug’s been have By construction visible. fractal- is (2004) tubular Narbonne’s anus and or 1994) (1973, mark mouth clear no a and crease, of most simple in a is gut but it a cases 2004), as Ivantsov interpreted 1996, The been (Jenkins decay. has genera or allied of growth axis arrested central localized be of equally areas could They (2002) gonads. Ivantsov & as oval Dzik by interpreted Indistinct been have margins well-defined organs. without also welts internal as interpretation annelid so with segments, incompatible all are also almost muscular across of run features evidence Gehling by as contraction interpreted ridges are in Such polyps. found or jellyfish more as fungi, lichens, rugae, look growth and these exaggerated like but margins 2002), well-defined Ivantsov lack & (Dzik Russia uua rnhn tutrs1–2m in mm 2 – 1 structures branching Tubular yJnis(96 g 4.2D), fig. (1996, Jenkins by Dickinsonia m )adinterspersed, and m) tal et Dickinsonia m ALCHERINGA :Pu 1973, Pflug m: 20) These (2005). . n plausibly and from Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 rnhd bqiosfietblrstruc- of fine-tubular tures Ubiquitous branched. lb(i.7)wt ueos‘ numerous large Seilacher of a with name is (informal 7A) these of (Fig. best slab The on slabs. impressions same resting of the basis and the trails on supposed invertebrate motile other or rings? fairy Dickinsonia decayed or rhizomorphs Trails, fungi lichens. like and more and animals, of histology the ilmt,tgte ihfour with costata together mat), bial (micro- surfaces ‘elephant-skin-textured’ on hanseni Phyllozoon rbbyagsecp tutr (Seilacher structure al escape et gas a probably howchini eo la mrsin of impressions Crisply clear below 1970). embedded ‘ deeply Martinsson defined more of cm are 2 (hypichnia and others thick mm wide, 2 – 1 1970). (exichnia) Martinsson moulds ‘ of (hypichnia Some of bed sole the the on imprints concave as preserved ‘ three lbi challenge. a Seilacher is slab 1992, al (Runnegar et gen- agreed are details erally topological These biomat. ‘ above originally h atndelpia rs-eto of cross-section more elliptical the ‘ flattened with The fossils these inflated of identi- worm (1995) fication Jenkins’s A by implied (1969). is burrow Glaessner by interpreted ALCHERINGA 05.B n fteeinterpretations, these of any neither By Seilacher 2005). by burrower worm hyaetolreadnto h aeplane same the on not and ‘ large too made are they have could slab Aulozoon Aulozoon ‘ Aulozoon 2005). . 03 05,btitrrtto fthis of interpretation but 2005), 2003, . Dickinsonia Aulozoon Dickinsonia aaohcstubularis Palaeophycus n single a and pig 99 h ato hc is which of last the 1949, Sprigg, Aulozoon Dickinsonia eealdee ntesa,so slab, the in deeper all were ’ a osdrdeiec faflat- a of evidence considered was ’ a enitrrtda worm a as interpreted been has Phyllozoon a aebe ri,as trail, a been have may ’ r ulrle sandstone full-relief are ’ Aulozoon u ohaoeand above both run ’ rtuigfo behind from protruding ekn eln 1978 Gehling & Jenkins nor Fg B r unlike are 6B) (Fig. Pseudorhizostomites and Phyllozoon Aulozoon n h surficial the and ’ Phyllozoon tal et Dickinsonia tal et al 1847). Hall, Dickinsonia 03 and 2003) . Aulozoon because ’ (2003, . nthis on The . are ’ EA OF DECAY huh ohv ie ihnsediment ‘ within If lived zoon 2002). Seilacher have & (Grazhdankin to thought nepeaini ht‘ that alternative is An impressions. interpretation fresh equally yeilrioop ietoeo modern ( of fungi those bootlace like rhizomorph mycelial rtalso ‘ of interpretations, thallus or ( these modern By Dickinsonia of 1997). ascolichens those Paradise like rhizines crustose system a lichen or 2005) of Bruhn & Mihail mycetes; oaino l four all of location one emntoso ‘ of terminations rounded adtn,nme fFg E n would and 1E) capping are Fig. of on markings 2 number (negatives faint sandstone, impressions these of raised speci- Fedonkin decayed Some variably of 2001, mens. be to Fedonkin marks appear 2002) & tool (Ivantsov or marks, off or trails vs spectacular show on this of not burrows does of case 7A) (Fig. in odds any specimen 1 even In (about one for alignment). rare chances a is 16 it coincidence, a hs eetal eann rmbefore (Seilacher from mat al et microbial remaining if the of preservation trails overgrowth crisp were the these and with worm, burrow coelo- mate a a of cross-section were circular nearly this more flat- if extreme required the tening sand, pushing biomatted flatworms are through problems for Other difficulties burrow. physical or trail the as ht‘ that loilsrtoso ungr19,Droser al 1992, et Runnegar of illustrations some also pen, sea a zoon as interpreted ally Seilacher of interpretation the follow age ihohrwnoe fossils winnowed other (Gehling with and biomat tangled the within like lived sediment it that (2005) Aulozoon te osl nepee stal,resting trails, as interpreted fossils Other 05.Iarewt Gehling with agree I 2005). . 05 ieinterpenetrated like 2005) . ntesa r negon(i.7;see 7A; (Fig. intergrown are slab the on ie ihntesdmn,te odid so then sediment, the within lived Aulozoon tal et ,bcuete r netie and intertwined are they because ’, Dickinsonia a aebe h mushroom the been have may Aulozoon DICKINSONIA 05.Atog convention- Although 2005). . Phyllozoon a oyfsi,adalso and fossil, body a was ’ rilramellea Armillaria Dickinsonia .Ti ssgetdby suggested is This ’. . ahrta being than rather , Aulozoon Aulozoon Xanthoparmelia tteedof end the at Dickinsonia tal et Pteridinium Basidio- : .I hsis this If ’. a a was ’ (2005) . Phyllo- Phyllo- tal et 231 ; . Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 ags o ndslya h ot utainMsu,soig23 showing Museum, Australian South the at display on now Ranges, interpretations. 3 RGR .RETALLACK J. GREGORY 232 rs ak,o akls htwould that swimming or creeping backfills, a of or motion indicate rims, marks, bulldozed brush undulation, trails lateral supposed single the of shows a None 1E). (Fig. of size different in slightly traces are they because grazing individual reat- or and also These displacements tachments successive found specimens. not are well-preserved halo around the marginal preserve but curiously 1E), filamentous yet internal Fig. lack impressions, and of rib effaced, 4 most sand- the and are these capping 3 numbers on sunken stone, are (positives markings faint impressions these of platform. a Others into sediment bind and surfaces, elephant-skin-textured on suppress produc- growth to creatures, microbial mucus flat large of such level for tion unusual an for call Seilacher (after four 7 Fig. iknoi costata Dickinsonia . A oto flresa rmteEicr ebro h anlyQatiei ahu og,Flinders Gorge, Bathtub in Quartzite Rawnsley the of Member Ediacara the from slab large of portion , tal et 03 05 Gehling 2005, 2003, . tlatoenro omtal n iepedmcoilmtwt lpatsi texture elephant-skin with mat microbial widespread and trail, worm narrow one least at , tal et 05.As hw r lentv nml( animal alternative are shown Also 2005). . esi rssgetv fatalcnas be also can trail a of speci- suggestive these arcs of in mens Arrangement outward center. grow a which from lichens, colony in the common of is necrosis central marginal with of lower rhizines preservation The the 3B). thick- (Fig. at measured ness and specimens, distinctness of decayed extreme as here xenophyophores. and lichens character- fungi, Such are of istic mouth. mechanisms a a feeding of than diffuse rather means foot, by digestive feeding and intermittently worm-like a for argued iul ako uhidctoso motion Gehling by of indi- noted indications also moving was such a outlines, of of Lack oval swath vidual. and continuous ghost a trails show not supposed marks these resting All organism. hs upsdtal r interpreted are trails supposed These hloonhanseni Phyllozoon ueos‘ numerous , Dickinsonia B n ihnfna ( lichen-fungal and ) tal et ALCHERINGA 20) who (2005), . Aulozoon moving sp., ’ C ) Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 ii et ak fprt,sdrt or (Wade siderite burial Gehling during 1999, pyrite, Gehling 1968, early of formed or masks calcite skeletons death mineral rigid other or agglutinate 1999). known Hughes angular cephala well & trilobite (Webster of so of compaction hint fracture, burial from a brittle there or shards is 1982, speci- 1992) Runnegar 1972, irregular Jenkins Wade and also In (see distorted xenophyophore. mens the or of lichen over- none by a substrate propagated of been growth have microbial also could irregular (Gehling an draped ridges bodies thin and be over to bumps thought been with have 2002). specimens (Fedonkin Other carapaces dorsal these chit- inous, that symmetrical, view bilaterally the coherent, were falsifies which wrinkles splits, as specimen or same the of to parts 1A). other untransferred a Fig. is in distortion gives localized (also Such shape subrectan- 1A) ellipsoidal to than gular (Fig. rather outline margins spindle-like of cases, than other In lobation wider 1D). (Fig. grown side one right-hand has the of 1D) of side (Fig. distortion left-hand a specimen The always symmetry. not is ideal this but slightly torn, and overfolded, marginally formed, Some biopolymer? or Mineralization 3C). Fig. com- example, almost (for on many rings bedding by plete discovered suggested single as of be planes, exposures will large that is suitably a model circles ring of fairy complete the fruiting of 2000). prediction peripheral (Ingold A mass to mycelial buried due central is complex arrange- ment more circular The to mushrooms. of circles swaths of simple These from alignment rings. fairy vary as common known generally the fungi, by explained ALCHERINGA ihSuhAsrla pcmn.Many found specimens. been Australian has South calcite with or siderite pyrite, hs bevtosfliyteie of idea the falsify observations These Dickinsonia tal et 05,btbmsadridges and bumps but 2005), . r rnldadde- and wrinkled are tal et 05.No 2005). . EA OF DECAY yrxdsbfr uil sdemonstrated as iron burial, of before precipitation hydroxides and encrustation micro- with bial comparable is fossils Ediacaran Earth in soft- time for other or any history. at fossils fossils Ediacaran bodied other for including invoked fossils, Neoproterozoic Dickinsonia only few is pyritic a for preservation of mask explanation (Retallack death the palaeosols Finally, by 2001). oxidation indicated atmospheric levels even (1999), Neoproterozoic Gehling in by sedi- of envisaged cm as 2 pre- – ment 1 to only beneath needed pyrite unrealistically be cipitate would in- An redoxocline biologically steep mineralization. pervasive of duced show but evidence 1994), no (Retallack been pruina have lichen may a which carbonate 1973), isolated (Pflug have crystals Namibia (Matten from plants fos- sils Ediaracan fossil Silica-permineralized of 1973). histology the study- in ing invaluable been a has section, that technique thick polished pyritized in to microscopically Heavily contrary 1E). relief, Russian pyritization (Fig. most by the observation then have effaced explanation, should most the were specimens characteristic masks the death Russian of conserves ribbing & than (Dzik obscures rather surface pyritization Thus, knobby 2002). nodules Ivantsov coarsely pyrite ellipsoidal a framboids thick with in it isolated oping of which ribbing nodules, the or either obliterate 2001) Reitner & as (Steiner fossils Ediacaran occurs Russian Pyrite with 1981). Dilcher associated & (Retallack smelly pyrite product of or weathering common yellow a jarosite, not with are fossils Australian Ediacaran or South pyrite. framboidal after knobby, textures tubular cubic not and but 6), felty (Fig. outcrop, smooth, patina are shows modern iron fossils the that in Ediacaran ferruginized Australian South h n eaii aiao many of patina hematitic fine The Dickinsonia n a o endemonstrated been not has and , DICKINSONIA Dickinsonia hudb examined be should Dickinsonia fpyritic If . envel- , 233 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 daaa osl rmascae small associated with from coeval been fossils have Ediacaran inverte- to thought infaunal brates, coelomate environmental for have and partitions shelter would marine food, environments shallow provided in intertidal fungi and or lichens Earth sessile three-dimensional Snowball Large dubbed 1992). (Kirschvink been have which and relief creating compaction. in burial resisting lignification to structural thin subordinate role that a but played in mask matrix, greatly death venation inorganic burial fine sandy before this of oxides impressions iron improved microns few fine a of with Dilcher Ferruginization & asso- 1984). (Crane than leaves were less ferruginous much fruits ciated burial Woody by venation. compacted and relief of thickness cuticle have strength their impressions to leaf proportional Dakota Dilcher These Cretaceous & 1981). (Retallack in the Kansas in of deposited Formation levees sandstones fluvial fossils ferruginized leaf angiosperm in for (1977) Spicer by iiain fa nmlwtotacir- a hand the on without If other 1982). animal (Runnegar system an culatory of as- limitations diffusional and oxygenation, requirements oxygen atmospheric suming on straints Dickinsonia implications change Global 3 RGR .RETALLACK J. GREGORY 234 03 a uigsqeta psdsof episodes (Hoffman sequential glaciation Schmidt during & global was Williams 1992, 2003) reg- Jenkins large, from 1992, to symmetrical fossils fossils ularly organized Ediacaran less di- small, evolutionary of and coincidence a versification appearance been the have not that effectively may It more (Retallack 2004). mats cooling microbial pre-existing global than and would in sequestration, lichens aided carbon under promoted formation have soil acceler- and produced ated prolonged have Deepened, would oxygen. it 1994), (Retallack a enue opaecon- place to used been has Dickinsonia Dickinsonia a lichen a was tal et (Fedonkin 1998), . oslebysi hshrts(Xiao phosphorites 2000). in embryos fossil (Seilacher burrows eas 1.4-m-long because Seilacher A 1993, 5). (Zhuravlev (Fig. interpretation jellyfish xenophyophoran fossil genuine than compac- tion and burial to resistance but stronger of on 1989), (Seilacher lack bilateral musculature and marginal obvious structure 1992) the quilt-like on (Valentine symmetry, only polyps not or founder 1956) Moore Interpretations 5). of (Fig. to compaction resistance strong burial and in- 1B), growth other (Fig. to dividuals reaction indeterminate allelopathic 4), – 3 isometric (Figs construction 6), fractal-tubular (Fig. by falsified is for Wade 1979) inter- 1968, Morris Conway worm Termier 1972, popular & The (Termier 5. pretation Table in biological bold various of for interpretations features Falsifying Conclusions yvru fterpyei eainhpwith relationship discussed. groups phyletic other their predictions of testable virtue generate by unfalsi- only (Seilacher remain but also clade fied, 2003) extinct Fedonkin completely 1992, Ediacarans a that Suggestions are C). – 7B swimming (Fig. leaping, lively, compared with seem 5), may it (Table unappealing unfalsified however remains fungal- model the lichen Only 1D). (Fig. skeleton agglutinative- mineral brittle an of than fragmentation or rather failure evidence deformation and variable plastic 6), highly for (Fig. 1A), diameter (Fig. microtube symmetry eral bilat- chlorophyte include of interpretation the xenophyophore features of falsifying those as seaweed too such uni- not siphoneous cells, though other with 1972), compared Tendal large cm: (25 xenophyophor- an living largest the than larger Dickinsonia Cladophora sjlys Hrigo & (Harrington jellyfish as tal et Dickinsonia tal et Dickinsonia Rnea 95.Key 1995). (Runnegar 05 sunlikely is 2005) . Dickinsonia 03 05 and 2005) 2003, . ALCHERINGA r hw in shown are Dickinsonia Dickinsonia a much was o a for tal et . Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 ALCHERINGA

Non-lichenized Xenophyopore Cnidarian Polychaete Annelid Turbellarian Lichen fungus protist Cnidarian Polyp jellyfish worm worm worm 100% below 100% below 81% below 81% below 75% below 69% below 69% below 63% below Areoles Lamellae Tubes Wrinkles Wrinkles Segments Segments Wrinkles Areolae around Lamellae around Tubular Mesenteries Mesenteries Segments Segments Wrinkles torqued midline midline alternate pseudo- pseudo- distorted by distorted by at midline growth bilateral bilateral muscles muscles Central stipe Central stipe Growth initial Mesenteries Mesenteries Gut Gut Gut pseudo- pseudo- bilateral bilateral Isidial initial Hymenial stipe Cell initial Mesenteric Mesenteric Pharygneal Pharygneal Pharygneal initial initial pouch pouch pouch Rhizines Rhizines Pseudopodia Tentacles Tentacles Parapodia Parapodia Parapodia Apothecia Peridioles Resting cysts Elliptical Elliptical Elliptical Elliptical Elliptical gonads gonads gonads gonads gonads Fairy ring Fairy ring Trail Trail Trail Trail Trail Trail Decayed thallus Decayed thallus Degraded test Resting trace Resting trace Resting trace Resting trace Resting trace Growth rugae Growth rugae Growth rugae Growth rugae Growth rugae Large intestinal Large intestinal Large caecae caecae intestinal caecae Podetia Scyphi Subordinate tubes Nephridia Nephridia Small intestinal Small intestinal Small intestinal unexplained caecae caecae caecae OF DECAY Up to 1.4 m long Up to 1.4 m long 1.4 m unlikely Up to 1.4 m Up to 1.4 m Up to 1.4 m Up to 1.4 m 1.4 m unlikely for unicell long long long long Isometric Isometric Isometric Isometric Isometric Isometric growth Isometric growth Isometric growth indetermin-ate indeterminate indeterminate indetermin-ate indetermin-ate unexplained unexplained unexplained growth growth growth growth growth

Areoles upper Lamellae upper Xenophyae Tentacles upper Umbrella pellicle Unifacial Unifacial Unifacial DICKINSONIA side only side only upper side only upper side structure structure structure side only unexplained unexplained unexplained Competitive Competitive Competitive Competitive Thickening Thickening Thickening Thickening reaction rim reaction rim reaction rim reaction rim unexplained unexplained unexplained unexplained Raised Raised Agglutinate Raised impression Raised impression Raised impression Raised impression Raised impression impression impression mineralized unexplained unexplained unexplained unexplained unexplained of crustose of crustose exoskeleton habit habit Compaction Compaction Agglutinate Compaction Compaction Compaction Compaction Compaction resistance due resistance due Mineralized resistance resistance resistance resistance resistance to chitin to chitin exoskeleton unexplained unexplained unexplained unexplained unexplained

Table 5. Biological models for features of Dickinsonia (with falsifying features shown in bold). 235 Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 oslrcr ffniadlces uhas such lichens, and the fungi with of but jelly- record cnidarians, bodied fossil soft and of worms, that fish, with not parable remarkably masks the of impressions death Instead, deep 1999). microbial (Gehling or or cementation (Seilacher agglutinative fracture of brittle skeletons suggestive of to evidence fragmentation any (contrary there decay Gehling 1989, Seilacher Neoproterozoic regime different of fundamentally a of idea the undermine palaeosols and fossils poraneous of decay decay of indications progressive These 2002). of Bottjer & (Dornbos evidence euxinic embryos clear fossil Neoproterozoic show phosphorites, in marine black Even Butterfield 2005). 1994, (Retallack fungi and 1990), h ahnm of 1994). taphonomy (Retallack walls The cell the fungal as of such biopolymer chitin tough a to due likely struc- ( fruiting tures actinobacterial (Retallack content fossil oxidized carbon 2001), organic highly low carbon yet organic isotopically red, with palaeosols includes evi- Neopro- the Other terozoic, during decay decay. aerobic for thickness aerobic dence in to variation attributable shows found, compaction. when burial to Dickinsonia resistant com- raised as pressions preservation is fungal- a affinity of lichen suggestive Especially rainforests tiered. to com- are mats all microbial from virtually munities and environments, inter- to tidal and marine plants shallow in algae, live protists, invertebrates of because variety affinities wide on light a shed Narbonne not do & 2002) (Clapham tiering community ele- (Jenkins habitats shallow startling to subtidal flat of Tidal create 1994). lichens (Retallack morels gance and and fungi as mushrooms simple organ- as even biological isms but 1996), of (Jenkins evocative complexity fossil complex and 3 RGR .RETALLACK J. GREGORY 236 Dickinsonia Primoflagella salclycmo osland, fossil common locally a is Dickinsonia sapesnl symmetrical pleasingly a is ooo Ivanovski & Sokolov : Dickinsonia Dickinsonia tal et tal et n te contem- other and tal et 05,early 2005), . 05.Nris Nor 2005). . 93 and 1983) . eemore were scom- is inr aito uigtelts Precam- 1994). latest (Retallack brian the during radiation evolu- tionary metazoan and change important global an for force been have would mushrooms fossils, the of Ediacaran obvious most As and toughest compaction. largest, burial the to resistant very and flexible, moderately ground-hugging, strate, Dickinsonia Jahren B B ites Thucomyces C B B B References discussion reviews. useful and much and for Erwin Jenkins Doug Richard Waggoner, Jim Ben Fedonkin, Gehling, Mischa and Hagadorn, Geology thank Whitey I of Adelaide. of Department University Geophysics, the casts. of collec- resin large tions the quality me showed Jenkins high Richard gene- supplied and Museum rously Australian South collections the to of access allowed Pledge Neville Acknowledgements UTTERFIELD RUTON RODO ERTRAND EAUMONT HALONER ihn fNrhAmerica North of Lichens e ae,C,xiii CT, Haven, New pigr eln 65-82. Berlin, Springer, aezi nteesenitro fNrhAmerica. North 7 of Tectonics interior eastern the the in of Paleozoic models numerical stratigraphy: and Orogeny abig nvriyPes abig,125-129. Cambridge, Press, University Cambridge 94 o-aclrln lnsfo h Devonian Ghana. the of from plants land Non-vascular 1974. A etArcnOoesadCru-tatcCorre- Circum-Atlantic and latives Orogens In African craton. West African West the of sequences Cover ..S Taxa Problematic of A.B. Significance the and Metazoa In traces. ‘medusoid’ fossil jellyfish with aebooy31 Paleobiology (Hallbauer FFATON .. S I.M., , .. 91 ec n aoaoyexperiments laboratory and Beach 1991. D.L., , tal et -S ,R.D.D .. M W.G., , . Q C., , IMONETTA ARFATI . 05 rbbePoeoocfungi. Proterozoic Probable 2005. N., , . T P., , , 03.Lk ug n lichens, and fungi Like 2003). . aaotlg 17 Palaeontology 389-416. , a rl tahdt t sub- its to attached firmly was HARNOFF Spongiophyton Aurelia UINLAN . M J., , ROMPETTE 165-182. , ALLMEYER tal et ENSAH &S.C ..&S & S.D. , Dickinsonia ,G.&H þ n oermrso some on remarks some and 97 ubr2001, Hueber 1977, . 9 pp. 795 ..&C & M.K. , ,R.&B OUSSINE aeUiest Press, University Yale . &J.P.L h al vlto of Evolution Early The 925-947. , ONWAY ALCHERINGA HARNOFF AMILTON and -P ELLION OUCHKINE ECORCHE M ihn or lichens RANE Prototax- ORRIS . 1988. J., , . 1991. Y., , . 2001. S., , M.D., , ´ eds, , ,A., eds., , The , Downloaded By: [University Of Oregon] At: 23:32 3 September 2007 D C C C D D C D C C ALCHERINGA C C C RAWFORD RANE ONWAY ONWAY ONAGHAN OLLINSON LOUD LAPHAM HERNS ROSER ROSER OYLE ORNBOS e ot ae ultn26 Bulletin Wales South New H oit fAeiaAsrcs34 Abstracts America of Society nu omto,suhetChina. southwest phosphatized Formation, Doush- putative Neoproterozoic antuo the of from biogenicity embryos sponge the of test d,Clmi nvriyPes e York, New Press, University 139-208. Columbia ed., 173-122. eds, oit fAeia35 picture? 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