7

The Ediacaran-Cambrian Transition: A Resource-Based Hypothesis

for the Pise and Fall of the Ediacara Biota
Alan j. Kaufman

ABSTRACT

Ediacaran oceano hosted a strange world of exotiç soft-bodied forma that fere a failed early evolutionary
experiment in nlacroscopic lide. These enigmatic oiganistlls appear suddenly in sedimentary rocks its old as
573Ma abole a glacial diamictite in Newfoundland, :utd, in most successioils,a profound and equally puzzling
negative carbon cycle ítnomaly known :ts tlle Shuram excui'sion. The Ediacarzul biota died out cear the
Ediacaran-Cambrian botmdary (<541 Ma) coincident with a second strong negativo õ''C excursion. From
Ediacaran aches,the ploli6eration of complex féeding trajes, soft-bodied artllropod tracks, and mineralized
skeletons in file succeeding Fortunian stage led to {t diverso landscitpe of modem phyla by the detonation of the
Canlbrian explosion around 529Nla. Tais chapter providos a review of the profound clxmges in the carbon,
sulfur, and strontium cycles acrossthis critical transition in arder lo better understand the redor history of the
oceano,as well as the tectonic, clinlatic, and biological eve1ltspieservedin its sedimentary archive,and further
proposes a novel resource-based llypothesis for the ride and rali of the Ediacaran brota.

7.1 . A BIOGEOCHEMICAL PERSPECTIVE the TI'ansgondwanan mountains may have dt'awn enough
CO, out of the atnlosphere to contributo to episodic
From çiimatic, geochelnical,and paleontological cooling of surfhce environillents }eading to {he developmen t
perspectivas. the Ediacaran-Cambrian tl'ansition stands of regional icesheetsand the deposition of glacial dianlic-
out as an interval of profound globo! chtulge pieserving [ites [ct, Rayrpio ef c//., 1988], ínc]uding the Gaskiers in
evidente in its sedimentary archive 6or {he coevolution Newfound[and, on at ]east eiylt pa]eocontinents [H(Záginír/i
of lide and environment (Fig. 7.1). Across this criticam a/z(/ 1,/, 2009]. The weathering flua of nutrieilts to the
[ransition, the initia] driver of oceanographic and oceanomay cave simiíí\rly contributed to global
biological evento is ellvisioned as tectonic in natura, cooling by stimulating primary productivity, theleby
related to widespread Pan-Africitn orogeny as east :tnd increasing the flua of organic carbon to the deep
west Gondwana fere sutured [Sgl/fre t'/ a/., 2006] in the oceano,as well as the buildup of oxygenin the
aftermatll of the ultimato Cryogenian ice age (aka snowballatmosphet'e,which is a phenomenon broadly known as
Earth event; K7rsc/r}/rlk, 1992; HoZ#lirr// e/ a/., 1998). the NeoproterozoicOxygenation Event (NOE) [Derry
Subsequent erosion of uplifted Ediaca ran terrains deliv- et a!., \992\ Des M«rciis et a{., \992.\ Ka!!ft7tattet {ll..
ered thick peles of sedimenta to the oceano, as weli as \ 993, \991\ Catttpbelt altd À!:eit. 2üQ8\ Har(!isto et at,,
bio-limiting nutriente, especially N, P, and Fe [Dtlry 2017; OcA a/id S/t/e/(Zx-Z/ro!{, 20 12].
e/ a/., 1992;Kííadilia/re/ a/.. 1993].Silicate weathering of In this review,the redox history of the oceailsacross
[he Ediacaran-Cambrian ti'ztnsition is considered in ]ig]lt
Department of Geology, Earth Systerll Science of tumultuous chances in the exogenic carbon, sulfur.
Interdiscipfinary Center, University of h4aryland, College Pack, ttnd strontium cycles;most notableale the pi-ofound
N4D,USA neg:ttive õ';C excursions that bracket the filst and last

Cllelltosttatigta!) t} Àcl'oss a..f(\lol'Clttoltological Botat(tarifas,Gemi)Itysic«!Ntottogrttplt 240, F\tst E.d\ttoh

Editei by Alcides N. Sial, Claudio Gaucher, Muthuvairítvasamy Ramkunlar, and VatderezPinto Ferreira
© 2019 the Anlerican Geophysícal Union. Ptlblished 2019 by Jolln Wiley & Sons, Inc.
115
116 CHEMOSTRAriGRAPHY ACROSS MAJOR CHRONOLOGiCAL BOUNDAREES

Precambrian Phanerazaic
matgrounds

%
Figure 7.1 The agronomic revolution or Cambrian substraterevolution clepictedin thís modlfiecl illustratio]] by
Peter Trusler reveals profoun(! chances in bioturbation across the Ediacaran (E)-Canlbrian (C) trarlsition. The
enigmatíc Edlacaranbioma(left) is rooted in ubiquitous microbíal mais that carpeteclthe seafloor (Precambrian
matgrounds) and largely sealedthe a noxíc sediments beneath from the free exchange of gases.The showdiffusion
of sulfate into these mais provided an oxidant for microbial SLilfatereduclion, leading to the buildu}) of toxic H,S,
a proposed criticamnutrient for the Ediacaran bioma.Thehorizontal burrows of animais within or beneath the mata
and the activities of the Ecliacaranorganismoon its surface, whtch a})pearvery late in the Ediacarangame, hacl
little effect on seclimentarylayering or the releaseof gasesto seawater. In contras{,the deep (tive of early Cambrían
(right) animais untothe sedimenta in search of foocl and shelter clisrupted the mais 'andallowed the free exchange
of gasesacros$ the seclíment-u,aterinterface, inclucling O,. Ventilation and mixing of the sedimenta is implicated
in the demite of {he Ediacaran brota, if H:S was a criticar physiological resource,as well as the builclup of sulfate
in the oceano. Reproduced with permission of PeterTrusler. (Seeinsereáorco/or represelltat;on of the #gure.J

appearanceof the enigmatic Ediacaran brota in post to more than a meter in length [Na/bon/ze, 2005]
successions.
To some theselarge and complexly orna- Understanding the position of these strange organismo,
mented organismo appear as saemgroup animais [e.g., typically preserved as three-dimensionar (3D) casta and
Rlillliegar, \982; Erwitt cf {l{.. 20\ \; Btld(! «n(t Jensett. molds in soft sandeand silosor as carbonaceous compres-
2017; Z)I'os'er a/ir7 Ge/l/íng, 201 5], while to others tllese sions in thin bituminous limestones [Wade, 1968;
soft-bodied tubular and fractally constructed fonns rep- G}'azhdallkitt. 2nQ4\ Nai'bot\tle, 2QQS\Fedottkitt et a!..
resent symbiotic lichens [Re/a//ack, 20 13] or as giant ulli- 2007], in the tme of lira is one of the greatestcurrent
cellular protists with no clear representatives in Cambrian cilallengesin p:tleobiology.
and younger smas (i.e., the vendobiont hypothesis; Rather than to take a ilaorphological appioach in plac-
Seí/rr('/iep',1989). Most modem worket's, hoxx'ever, regard ing ornamenta of the Ediacaran bioma on this trem, it
the Ediacaran biomaas representativeof a multitude of seems worthwhile to apply a biogeochemical perspective
multice:pular phylogenies united by their soft bodies :\nd and ask Hirst whether these organisnls might cave had a
largo size. common metabolic stiategy. For example, osmotrophy
Ediacaran-type fossils (which base been !eported at has beenconsideredviable in a dissolvedoigailic carbon
nearly 30 1ocalitieson 5 continente)are typically in the (DOC)-rich ocean [Ro//u/ian e/ aZ, 2003; Z,a/7a/}1/}7eer a]] ,
centimete] to decimetei'l-ande,with somegiants ranging 2009] given the pt'esenceof a largo surEaceárea presented
 THEEDIACARAN-CAMBRIANTRANSITION
117

by the tubtitar and fraçtal morphologiesof the Ediacaran Ediacaran game; it htts beensuggestedto play a role in
brota. ExEendedsurface are:ts could allow for the simple the demise of the Ediacaran bioma[e.g.,l.(Wzí/n/?re e/ a/.,
diRusion of dissolved nonpolar organic molecules(and 2013; Bz/(/í/a/i(r Jc.n.çe/z,
2017]. The oldest potential yet
oxygen) througll the thin walls of the organismo in ordem' problematic burrows (attributed as .4/c/raeonasõ'a)
ale
lo fuGI tlleir metabol ic activitíes, a 'ózí/nfngthat these wel'e reported stop -565Ma deepwater turbidites in
respiratory in natura. Osmotrophic üeedingis an intriguing NeMoundland [Z.üi e/ a/., 2010, 2014], but simple,
hypothesis,especially considering that inany re!)resen- unbranched, and horizontal surface tFacesreliably attrib-
tative taxa lived below tule photic zona [ÀrarZ)o/i/ie uted to bilaterians appear abundantly in sedimentary
and Áiíken, \99S; Dah'yfnple attd Narbot21}e, \996; rocks around 550 Ma [Jt'/!se/z, 2003]. T1lese simple bur-
i\ acNattglitotlet al.. 2QQ0:Clapllan}ef a!., 2QQ3;!'voos roxx'ing trajes (e.g., He/rn//ir/rofc/fc/?/rífes) are believed to
e/ a/.. 2003] and that theie is scant evidente for any of represent the peristaltic movement of animais through
these organisms llaving a mouti], gut, or ânus. Notably, in the lbict'obial mata at or very near to the sediment-bater
modem ecosystems, only microscopic bactéria with largo interface. Recently, more complex surface trackways of
surfaceáreasreiative to total volume are abreto survive bilaterian aninlals with paired appendagesllave been
by osnlotrophy.
If the organicmembranas
of the reported alongside simple burrows in termineil Ediacaran
Ediacatan biota (inclltding both tubular erniettalllorphs strata of SoutoChina [C7lí?/l
e/ a/., 2018].Neai the
and fracta! rangeomorphs) were thin and thcir interiors Ediacaran-Cambrian boundaiy, icllnoüossils become
fere hollow or band nilled. then their surface área to more 3D with evidente of a deeper deveunto sediments
volume would appi'oach that of the osmotrophic bactéria while probing or farming (agrichnia) 6or food [A/ringrrno
[Zid7u/pz/}ie
ef a/. , 2009], allowing for this feeding strategy. e/ a/., 2012; Z,a/rzgef a/., 20181.By piercing and venti-
lince a large standing DOC pool was probably a long- lating Ediacaran matgrounds [Se//ac'#er a/ií/ P7/zlger.
stailding feature of the Neoproterozoic oceans (in con- 1994; Bo/{/er e/ a/.. 2000; Gougeo/ze/ a/., 2018; Ha/z/ioo
[rast to the dominant dissolved inorganiç carbon (D]C) e/ a/., 2018], Fortunian (the iatiHiedbasal stageof the
pool in modem seaways),the Ediacaran brota appear to Catllbrian period; Penaef a/., 2012)bioturbation would
have evolved largely associated with the ride of oxygen in h:we depleted the hydrogen suiütdelesotirçe requimd 6or
surface environments [cf., ?VlJr.sa//. 1959; Berk/le/ c//id the proposed Ediacaran metabolism and may have
&/a/-.ç/ra//,1965] and may have seen their demise with fala ultimately resulted in [he Cambrian buildup of oceanic
of the breathing gas near the Ediacaran-Cambrian sulf'ate]Ca/Üie/d and Earqu/ral', 2009]
boundary [Kfnzrr/a aní/ 14/ararzr7be,
200] ; Hnlr/ror ef a/., Tule evento described ]lelEin are viewed tht'ough a 50
2QQ3; Schtõ({erand G}'ot=inger,2001\ Wille el a1.,2008; tnillion year window spamüngthe presentlyundefined
cf a/. , 2013; Z/l«/zg e/ a/., 2018] . Following tais terminal Ediacaran stage(TES) [Xfao e/ a11,2016] and
Z,{{/7a//7/ne
logic, oxygen would have been one of the key metabolic thraugh the base\lCalllbritut Fortunianstage.One lide is
resoltrces6or the Ediacaran brota, whether they were ani- h'amed by the Shuram excul'sion (SE), an unpl'ecedented
maisor not. Given the inÊtunal and epihunal lifestyle of and highly debated alkalinity and negativo carbon cycle
tllany of these large pneumatic organismo, } explore anomaly pl'eservedin a variety of carbonatohcies on
whethe] hydrogen sutHide (H,S) produced below and multiple continente [X)iai/r/r a/r(/ .Ke/r/iedy, 2009;
within the ubiquitous microbial mataby sulfato-t'educing Grofzingeret a1.,20\ \; Scltr«get a1. 2Q\3; Cui et {tl..
miçrobeswould cave been tuleother critieal metabolic 2017lj xvhile on the other lide is the Cambrian explosion
resourceand further that the earliest form of bioturba- of lnodern animal phyla [.Ro:anal'ef a/., 1969].Within
tion was microbial growth unto the sedimentain order to this narrou teinpolal window, lidegot big rival'brlnnean(/
accesstheçorrosive, toxic, flammable, and smellyhydride Ge/z//ng,2003], smart [Cczróonea/i(/ Nai'Zloíine,2014], and
gas.Ratherthan tllaking the a priori assuillptionthat hard [Gel'/ns,1972; Gra/zr,1990;GT'orz]/zger
e/ í//. 2000;
[heseorganismoweie animais and somehow h:td to pro- Pnp'rer, 2007, 20 10; Z/il/ra/t l a/í(/ mood, 20081
tect themselvesfrolll toxiç H,S in a symbiottc relationship
jcf., Z)zeáol'
a/zd A/c'//roy, 20 } 6], 1 hypothesize an alternativo 7.2. CHEMOSTRATIGRAPHY
bacterial physiology]GP'í/zA(/r//ik/n {r/z(/ Gel'cüs, 2007] for
[he Ediacaran brota based on thioautotrophy (i.e., sul6ide The profusion of chemostratigraphic studies of shallow
oxidation in the absenceof light) that could explain their marine sedimentary successions that accumulated acmss
morpllological disparity. as well as their spatial tutd [he Ediacaran-Cambrian transition is testament to the
temporal distribution in terminal Ediacarall seaways. importance of tllis nlost çritical tl'ansitionin Earth'shis-
Bioturbation, or literally the bunowing and stirring of tory, as well as the semanal isotopiç researçh of BÍ// Ho/sel'
soft sedimenta by monte triploblastic (bilaterally tHoiser alKi K«play\. \9G6\ play!)oo! et al.. \980\, ÀÍanfrecl
symmetricwith a trufamouth, gut, and {tnus) anilntils in Sc-/ií(//após/ e/ a/. [1975], and Jzí/l Heücl' [l'?f:úr an(/
search of food aild shelter, appears very late in the Cbnpõron, 1976; Helzera/i(/ Ho(yk, 1976].These scientists
]18 CHEMOSTRATIGRAPHY ACROSS MAJOR CHRONOLOGICAL BOUNDARIES

were among tuleHlrst to identify largo swings in the cztrbon, and earlier Ediacaran [Kno/re/ a/., 2004]that set the stage 6ol'
sulfur, and strontium isotopic colnposition of seawater the origin of the Ediacai':tn brota and of animal& Foremost
proxies preserved in Precambrian and Phanei'ozoic basins. among the carbon isotope eventoin this tumultuous interval
The dynamic and expanding landscape of Ediacaran and is the enigmatic Shuram excursion (henceforth abbrevi-
Cambrian chemostl'atigraphictesearçhover the post 30+ ated as SE). The origin of this biogeochemiçalanomaly
years owes much to these early isotopic explorerx and its relationship to the Gaskiers ice age is one of the
While current studiesprovide rigorous sequentestrati- greatestoutstanding problemaof Ediacaran Earth history
graphic, biostrittigiaphic, iutd radiometric {trchiteçtures [Xfao ef a/. 2016]. First recognizedin middle Ediacaran
on which to gang chemostratigraphicdata, most are still strata fronl Oman [Blír/r.\ a/id /Varrer, 1993] and Soutl}
hindered by incompleteness of the sedinlentary record. Australia EPe//e/ a1, 1993],the deepstrittigraphicdivide
the absence of key lithologies (including interbedded çx'assoon found on other continents preserving I'emark-
volcanic aches), and/or diagenetic insulta. The largo ably similar õ''C trends [Gí'ofzirzgel'
ef aZ, 2011]. With
number of publications purporting composite refer- nttdir valuesdlttmatically lowel' than mantleinputs (ca
ente sections is testament to {he general acceptance of --5%o),its primary nature haslona beenquestioned
isotope stratigraphy as a critica] too] in the Ediacaran While the Gz\skiersdianlictite is remarkably well con-
and Cambrian periods. It is algo a manifestation of the strained by ractiometric datem [Plr ef aZ, 2016; see
ambiguity inherent in projecting local systemsoito the discussion below], the SE is not. In Ediacaran strata of
worjd stage[Cbl'será/ alzdKaud)piam, 2003].Intrabasinal Namibia, the biogeochemical anomaly is preserved in
correlation remains a critical aspect of reconstructing carbonato hcies of the Kanies and Mitra members of the
historical evento in deep time and is best served as an Kuibis Subgroup [ IHor)c/
e/ a/., 2015], whicl} sit hundreds
integrated dish. Technological advancemenÍsover the post of meters belos' an -547 Ma U-Pb zircon age çonstraint
25 years haveallowed for a proliferation of novel isotope [Gí'í?rzúiger ef a/., 1995; recalculated in Bola r/ng el a/. .
measurements(e.g.. Cr isotopes in carbonates; FI'e/ e/ a/., 2007; ]Varbo/l/re e/ a/. 2012] but provide a minimum
2011) that purport great insigllt to environmental cl)ange, upper bound. Closer to the event, many researcherspin
but theseI'equil'efurther caliblation, as well as strati- [he top of the SE to the 551Ma Doushantuo-Dengying
graphic and diagenetic teses[/íoo(ref a/., 2018] of whether boundary age in South China [Condo/i ef a/., 2005], but
they reílect local perturbations or global pÍlenonlenon. ullceltainty as to u,hether tlle Miaohe volcaniç ash that
The EARTHTIME initiative has accelerated advances in nes xMthinthe fiuuedtagerstãtte
belongsto the
geochronology lince 2003by providing calibrated solutions [)oushantuo or the Dengying is criticai]Kaz/chia/z, 200S].
to U-Pb zircon labor:ttories worldwide in arder to compare A recent study indicated thal the ttsh nesabovea notable
standard results, to minimize boas,and to improve absoluto sequente botmdary [H/z e/ a/., 2015], such that the upper-
calibration. Based on these enorts, sub-mi1lion year uncer- nlost interval of the Doushantuo (Member IV) could be
tainties on U-Pb zircon dares 6or both Ediacaran and muco older than it currently appears in most cllemostrati-
Cambrian periods are more and more coirulaon, and in {um, grapltic compilations [seeX/ao t r a/., 2016]. The onset of
we understand more about rales and the driving 6orcesfor the Shuram anomaly, which is coincident with tl\e global
biological and oceanographic chance. Applícation of these increase in seawatel alkalinity as evidenced by the sudden
ages,howevet,must still be tied to the stratigraphic record appearance of thick-bedded calbonates in otherwise
with tule understanding that most successionurepresent siliciclastic dominated successions,is equally uncertain.
mbsing time {tnd t1latçryptic unconformities or miscorrela- In the absence of dateable volcanic aches, three non-
tions can confound even the most preciw dares Tbe zircon-bttsed radiometric techniques nave been atteinpted
development of laser ablation U-Pb zircon techniques is a to constrain Doushantuo Member IV Using Lu-Hf and
tremendousadvanceallowing for rapid agecharacteilzation Pb-Pb techniques Ol} phospllorites some20m beiow tule
of bota detíital and vc)lcanicgrains. However,if this tech- unit, l?aiyb(/ e/ a/. [2002] documented ages of 584t26
nique is used to pre-characterizevolcanic zircons, tulelower and 599.3t4.2Ma, respectively.
A similar age of
piecision age distribution of a// grains determined by the 595 t 22 Mt\ was recently reported from Re-Os analysis of
rapid technique should be pubiished alongside the higher blaçk shale from the base of Member IV [Z/nr e/ a/.,
precisão:l ages of grains selected spwifically 6or chemical 20131;the sigilinjcant uncertainty and high MSWD wíts
abrasion single-ciystal analysen suggested to result fronl temporal chances in the
187Os/ossos
of seawater, se [t subset of the analyses were
7.2.1 . Shuram Excursion used to estimate a depositional age of 591t3Ma
Accepting any of theseunconventional radiometric con-
This criticam transition chapter focuses solely on the straints, the beginning of the SE in Souto China couid
TES and Fortunian intervala wllile acknowledging the Hall be çoinçident wiEh the end of the Gaskiers ice age oi
coupled climatic and geochemical events of the Cryogen ian even be somewhat older.
 THE EDIACARAN-CAMBRIAN TRANSITION 119

With {heseage uncertaintiesin mind, there are a .12-8-4 0 4 8 12

number of possible historical atternatives to consider. In
[he accompanying ct\rbon isotope coma)i]ation (Fig. 7.2),
the SE is shown to begin immediately abovethe 580Ma
Gaskiers diamictite (see correlation 2 in Xlao ef a/.
(2016»,suchthat all of the Ediacaian biomas
lie abole the
biogeochemical anomaly. Thus far. i{ is only in north-
western Cartada, w-here a putative SE has been identi6led
Ü
in carbonatestrata assig1ledto the Ganletrail Formation
in the \Verneckeand Ogijvie mountains [Jo/r/is/one/ a/.,
2013; À/ac-(/a/za/(/er a/. , 2013], that deepwater Ediacaran
organismare interpreted to llave lived be6olethe inferred
oxidation event. This view, based on proposed,but not
unambiguous,interbasinal cora'elations,is complicated
by the absenceof an SE in the Mackenzie Mountains w;-»-"l
where a depaupefate but recognizable assemblageof {

Ediacaran fossils (including complex and radially sym-

metrical disco and fronds, but no segmented forma) cave
previously been documented in rapperSheepbedstrata
belos the Gametrail Formatio!}. Previouscllemostrati-
graphiç research in the Mackenzie Mountains [KatfÓPlrrr/i ?

e/ a/., ]997] identiHledtwo strong positive õi3Cexcursions

in tuleSheepbedFormation [the upper interval now char-
acterized as the Jul)e beds, which contam the Ediacaran
fossi[s; À/r7cdona/(/ ef rr/., 20]3]. A]thotlgh carbonate
lithologies are lacking, it is plausible that the negativoSE
nes between [hese events, which wou]d support the view
entertained hera that all of the soft-bodied lide fol-ms
postdatedthe biogeochetnicalanomttly. To support the
alternativo conchision. Ediacttran fossils would need to
be discovered in the se-ct\lied June beds in the Wernecke
:tndOgilvie mountain$ or a credible explaníttion far the .12-8-4 0 4 8 12
:tbsence
of the SE in the thick Gametiail carbonatesof
õnC(%.,VPOB)
the Mackenzie Mountains must be presented.Insofar as
these remoto successions in northern Canada may hold Figure 7.2 Generalized trend in the carbon isotope composi-
the key to our understanding of Ediaçaran Earth history, tior] of marine carbonates through the TES and Fortunian stage
further research is warranted. of the Cambrian period. This compilatíon is !)asecfon cintafrom
Controversies surrounding the SE and its potential a wicle arrayof sourcesfrom successíons
in Índia IKaufman
correlativosinclude (i) its origin: wllether the event ef a/., 200ól, Morocco [/Wa/oofet a/., 200S], Nam]bia ]Kauámarl
representa tt lona-iived distui'banco of the global ocean et a/., 1991; Say/oret a/., 1998; Woodef a/., 2015], South
China [A4cf.a(/c/enet a/., 2008; Cui et a/., 2015, 201 6a, 201 ób,
DIC reservoir due to the oxidation of organic c:ubon
201 7], amar Ifike et .l/, 200ól, Índia IKauÁmanet a/., 2006;
or methane[Ro//i//zcln
ef a/.. 2003;Ffkeef a/., 2006; Tetvar/ and S;a/, 20071, Siberla IKno// et a/., 1995a; KauÉnTan
Kattj'll\atl et at.. 2QQl\ Bristow atld Ketltedy, 2nQ8, ef a/,, 1996; A4a/oo/ef a/., 2010 and referencestherein; Cu/
McFac!(!cliet a1., 2008; Bjert' 1»1clnd Cailfieid, 20\ \; et a/, 201 6c], South Australia [Husso/l er a/., 201 5], and the
Hrrsó'one/ a/-, 2015], (ii) reflecte conditions coilducive United States[Córserf/anc/ Kauf/7}an,2003; Hebert et a/.,
to authígeniç çarbonate precipitation [it/ac'(/o/ia/d 2010; Verde/ et .]/., 2011]. This compilation should be com-
ef a/.. 2013; Sc/n'írger a/., 2013; Cuí ef a/., 2017], or (iii) pared against those presented in Xfao ef a/. j201 6} insofar as
results from meteoric or burial diagenesis [KPíazff/za/l{/ the t.ipperreachesof the Shuramexcursion are not pinned to
Kcnnedv, 2009; Z)erly, 2010]. While we have docu- the 551Ma U-Pb agea{ the Doushantuo-Dengying
contact
mented clear petrographic and isotopic evidente for (see text for explanation). Posítionsfor ice agesare markedby
Ihe presente of authigenic carbonato nodules derived AAÂ. Note the uncertainties of the positions of the baseof the
from the anaerobic oxidation of methane associated TES anelthe end of the Fortunian. which rebateto issuesof cor-
relating cherllostraligraphic an(f biostratigraphlc events l)et-
with the SE in Souto China ]Clíí e/ a/., 2017]i\nd w,een basins.
Nanlibia, my personal bias is that this was a primary
120 CHEMOSTRA[EGRAPHY ACROSS MAJOR CHRONOLOGICAL BOUNDARIES

oceanogrtlphic pllenomenon related to the progressivo ocetulic sulhte is provided by time-seriesaní\lysesof

ventilation of the ttnoxic pre-Shuram oceano. strontiuill (Fig. 7.3; indicating a signiHicantincrease in [he
[n this view, the sudden increase in ''C-dep]eted cjelivery of radiogenic *'Sr to the oceans)tuld sulfur iso-
tlkalinity wits a direct result of the delivery of nutriente tope abundances[e.g., /'fke e/ a/., 2006; Katt#nan ef a/.,
lnd sulfato during intenso weathering of uplifted 2007; A/c.fü(/r/encf a/., 2008] during the unprecedented
Trítnsgondwananten'ains. These continental fl\!xes carbon cycle perturbation. In fact, given the remaikable
would have stilnulatedanaerobicmia'obial sulfato coupling of carbon, sulfur, and stro1ltiumisotope chance
reduction(MSR) throughout the oceanoand the u-ater during the event, either the SE representaa diagenetic
column production of carbonato u ith strongly negative conspiracy, or it is an indicador of truly global environ-
6t3Ccompositiona The sulÊttereducerswould havedined mental change [Ha/t'ers'o/i e/ a/., 2007; J(rzu!/han ef a/.,
on the abundant DOC (eventually depleting tais resource) 2007; Lee e/ a/., 2015; Cuí ef a/., 2017].
allowing for the oceano to become increasingly oxygen- Accepting tllis holistic scenario, by the end of the SE,
ated[lb/cEa(/(/en e/ a/. , 2008; Oc'/i «/l(/ S/ífe/íl3'-Z/roer, 201 2], shítlloxv oceanwater would cave been oxygenated enough
thereby driving the anaerobes unto the sedimenta by the [o stimu]ate the evo]ution and diversif]cation of the
end of the biogeochemical anomaly. This scenario, Ediacaran brota. These organismo appear very near to
inçluding the progressivesensoof ocean oxygenation, is the crossover
froin stronglynegativo-to-positivo
õ''C
supportedby recent U and Mo isotope measuwmentsof compositions. with the earliest example perhaps being
Shuram equivalent carbonates in the Doushantuo of Pa/aeopasc/r/r/nfs; most modem workers regard tais
Soutjl China [Kendír//c'fa/., 2015]. as xx,ellas from iodine Ediacaran form as a serially repeating body 6ossil [e.g.,
abundante measurementsof carbonates t'rom the Shuram Ha//ies, 2000; ,4nrc/{Égk( r a/, 201 1],although it has alter-
Formation of Oman [Har(/is/y e/ a/., 20 17]. Evidente for natively been interpreted as the earliest evidente R)r
intensoweathering of the continente and the buildup of bioturbation [Rogot'et a/., 2012]. The fóssil occurs in

0.7100 0.7100

e Doushantuo/Dengylng,Jiulongwan
o Doushantuo.Yangjíaping
O Doushantuo. Zhongling
0.7095
A Shuram. Oman
O Wonoka, South Australia
O Chenchinskaya, Southern Saberia 0.7090
B Khatyspyt. Northern Slberla

\ 0.7085
e
+
M 0.7080

0.7075 0.7075
Shuram excunion

!
0.7070
645 635 625 615 595 555 545 535

Age (Ma)

Figure 7.3 Generalízed trend in the strontium isotope composition oí well-preserred high-Sr marine limestones
through the Ediacaran period modified from X]ao et a/. 12016] with age constraints based on their "correlation 2"
and assun)ing the Shuram excursion is pinned to the 551 Ma age for the Doushantuo-Dengying boundary (see
discussion in text and Fig. 7.2 ca})tíon). Tais compilation is based on (lata from successions in Souto China l5a t,ak/
ef a/., 201 0; Cuí et a1, 201 51,Oman [Burns ef a/., ] 994], South Austra]ia ]Ca/ver,20001,southern Saberia[A4e/ezhfk
et a/., 20091, anel northern Siberia tCui ef a/., 201 6cl. The trenó marked by the thick gray lhe indlcates a plateau
of ca. 0.7080 followecl t)y a profouncl ride in "'Sr/mSrvalues coincident with the Shuram excursion up to as hígh as
0.7090 anel likely associatedwith intense weathering up uplifted terrains worlclwide. The trend then decliiles pack
to -0.7080 very near to the E(fiacaran-Cambrianbounclary. (SeeinsereÁorco/or /eplesentaf;on of the í;gt/re.)
 THEEDIACARAN-CAMBRIANTRANSITION
121

Unir 8 of the Wonoka Fbrmation of Souto Australia minerais se they are a mole direct proxy of seztwater
IHaines. 2000ljust as the c:trbon isotope trend :\pproaches conditions. T1leir relativo enrichments in Ediact\ran {tnd
[he0%«v:t]ue[/7i/.\.\'a/ief a/.. 2015]. Simi]ar]y, in Nanlibia, Cambrian successions in comparison with older strata
the llrst of the soft-bodied Numa Assemblage appears in stlpport the view of rising oxygen in surfaceenviron-
sandstoneof the Kliphoek and Aar members[Hr7//ef a/., ments, although nono of thesepaleoproxies are particu-
2013], inçluding Eln/erra, P/er/dlííftl/n, and Ralzgía, [ar[y wei} ca]ibiated. ]ron speciation measurements
immediately above the SE p!'eserved in the Kanies and (wlaiçh are better calibrated in shale from modem oxiç,
Maia lllembers. 6erruginou$ and su16idicenviromnents; Poli//on alia
Can!/ie/r/,2005) of fossiliferous Ediacaran sedimentary
7.2.2. Neoproterozoic Oxidation Event (NOE) rocks in Newfoundlandare consistentwith the post-
Gaskiers ride of deepwater oxygen that 6osteredthe evo-
Due to the desert climate. {he geological exposuresin lution and diversiHlcation of macroscopic life [CarÚie/d
southern Namibia are spectacular, and the views are e/ í//., 2007, but see Spér///ig ef ír/. [2015] 6or a contr:try
expansivo.In terms of an origin story, it is fere where view]. While theseanatysesplay a supporting role in our
time-seriesEdiacarancarbon and strontium isotope understanding of the NOE, the iron speciation proxy is
researchwas 6irst hatched [Kaüdina/i er a/., 1991, 1993] on consideredby l)}anyto caveonly local signinicance;resulta
simples graciously donated by Gerard Germe. Wulf fiam one basin to the next seen]fairly clearly Liedto the
Hegenberger, and Andy Knoll, along with straligraphic organic carbon and pyrite contents of the preseived
assistancefrom Charlie HoQmann who has played a last- s1lales in each basin.
ing role for a llost of internationalinvestigatorsever Temporal changesin the sulfur cycle recorded in the
lince. Frolll the baseof the Nama Group, one can see isotopic compositions of sedimentary sulfatos, car-
Ediacaran history play out as the carbonate }ithologies bonato-associated sulfato (CAS), and pyrite may also
becomeincreasinglydarker in colar from the Mttnt to the provido imporEttnt constraints on Ediacal'an and
Mooifantein reflecting their organic carbon contenta Cambrian redor conditions. Relativo to measurements in
[Say/OI'ef a/., 1998].Associated with the tonal transition older intervala, Ediacalan successions document :\
are dramatic increases in the i3C and 87Srabundantes of signiniçant change in the magnitude of fractionation bet-
well-preserved limestones (Figa. 7.2 and 7.3) {tnd the evo- ween reduced (pyrite) and oxidized (stilfate or CAS)
lution of mactoscopic lide. The isotope chances preserved phases[CalÜie/c/
rz/zdT/ia/lr(/r'ilp,1994;(ki/l!/ie/c/,1998]
in carbonates are believed to l-eflect proportionally This redistribution of sulfur isotopes has been interpieted
greateiburial of '2C-richorganic manterprompted by the to ieflect theonset of oxidative processes,includíng sulfur
coincidenteof high primary productivity and enhanced disproportionation. In dispropoJ'tionation, sulfur is recy-
ratos of continental weathering. The Ediacar:tn Sr iso- cled via both reductiveand oxidative pathways.On the
lope record illustrated in Figure 7.3 is consistent with reductive lide, the magnitude of kinetic sulfur isotope
Pan-African uplift of Himalayan-scale mountains and fractionation correlatesdirectly witl} extracellularsulfate
the related erosional stock to the oceano [.4.\'/nel'o/lref ír/., concentrations. Experimenta frota puro cultores of sul-
199 \ ; Der)' al:({ Fraltce-liinord, \ 996; Sq11ii'e {-f a!., 2QQ6; fate reducers indicate maxinla! fractionation of 66%o at
C«lll/)óc// an(f ,4//eiz,2008]. Insofar as there is a linear sulfato concentrations similar to inodern seawater a{
I'elationship between sedimentation rate and organic 28mM [S/p?z e/ a]; 2011], whi]e fractionations may be
carbon burial [Z)el'/' e/ at. 1992], atmosphericO. wi]] suppressed at very low sulfato abundantes (<200pM)
increase during orogenia evento but only if the buried [Haófc/ir e/ a/. , 2002]. On the oxidative sêde, the su]Hide
organic mass (along with reduced iron and sulfür) exceeds pioduced through MSR is typically reoxidized to ele-
that of the mass of organic carboi} el'oded[Kald;nan mentar sulfur, which is subsequently disproportionated
ef a/., 1993; Ca/npbe// rznd .4//en, 2008]. Geochemical
to sulfaLeand sulfide, by coupling with the reduction
modelaof secular trends in çarbon. strontium, and neo- of O:, NO.', or iron and manganese compounds.
dymiun} isotope changes through the Ediacaran imply Disproportionation reactions can thus augment the frac-
that most of the O, in the present atmosphere(ca. 21%) tionations induced during MSR, resulting in isotopic
could cave been generated [see algo Z)es lb/cn-a/sef a/., contrasts betweenreactant sulfato and product sulfide of
1992] during the NOE [Oc/i a/i(/ S/ífe/(/i.-Z/ion,2012]. >70%o. Interpreting the environmental signinlçance of
An independentbaiometer 6or Ediacaran oxidation stratigraphiç variations in the magnitude of fraction-
may come from elemental redox proxies, including Mo, V. ation, however, requiKS the recognition that the õ34Ssig-
:tnd U [S(orr ef a/., 2008; Oc'/l and S/i/e/ds-Z/iou, 2012; natures of CAS and pyrite may have beeninherited from
Prrr/frzef a/-, 2013] abundantes in black sltale, as \x,ellas diüerent pares of the depositional basin [Cu/ ef í//.
Ce çoncentrations in carbon:ices [14'h//ace e/ a/., 2017]. 2016a]. Sulfate incorpolation indo prim:try carbonato
Theseelenlents cave very minou concentrations in detrital sedimenta would occur within the water column, whereas
122 CHEMOSTRATIGRAPHY ACROSS MAIOR CHRONOLOGÊCAL BOUNDARIES

pyrite would 6onll either in euxinic botton] wtttersor 2013]. The origin of the "S-enhanced pyrites }algely
uithin sedimenta.Considering this sp:ttial separation, remítins :t mystery but is consistent with rapid ratosof
local sulfato availabiiity could dictate the sulfur isotopic MSR in the sedíments(or in anoxic bottom waters) stüll-
diHereilcebetweenCAS and pyrite, particularly if pyrite ulated by the presente of abunditnt nutrients :tnd organic
is formed in non-bioturbated and microbially sealedsed- substratos.More conceining, however,is the likelillood
imenta whele the water-sediment interface representa zt [hat previous iesearchershave underestimated the non-
signinicantdiüusion barrier [Bor{/er e/ íí/., 2000; /i]ke structurajly bound sulfur compoilents in these I'ocks by
e/a1, 2009]. using incomplete leaching teclmiques [cf.. A/ül'e/lco e/ ír/.
Evidenced by the widespread deposition of marine 2008; Tos/et'f/zef a/., 2017], resulting in higher CAS abull-
evaporites near the end of the Ediacaran in Oman and dances and lower ÕJ+Svt\lhes for their seara,ater sulÊate
elsewhere, some sulfur isotope studies conclude that the proxies.
overall sulÊate coilcentration of the oceano increased
from the SE to the Ediacaran-Canlbrian boundary 7.2.3. Basal Cambrian Excursion (BACE)
[Hltl'rgen ef «!., 200S;File et a{., 2006 Ha]versotta td
Hríirgen, 2007; 14/íre/ a/.. 2015].Tais interpretation is Carbon isotope trends !eading up to the Ediacaran-
consistent with the oxidation of pyrite in exposed Cambrian boundary are characterized by a plateau of
continental rocks during Pan-African orogeny, i)ut it inoderately positivo õ':C compositions that in some
could alternatively signify the releaseof sulfide from regíons hll through {he origin [Pt./er'/zaíy e/ a/., 1996;
sealedmats (and its conversion to sulfate via microbial Slli/f/l ef a/., 2016] before a pLmctuated negative carbon
sulflde oxidation) associated with the onset of bioturba- isotope event known as the basal Cambrian excursion
tionlCb/!#e/U í d Ãarzgzr/ral'. 2009; FHuef íz/., 2015: (BACE). This profound biogeochemical anomaly rivais
Ha/if.sooe/ a11,2018]. Enhancedsulfide oxidation pro- the SE in its depth, if not its stratigraphic throw (Fig. 7.2).
vides an alternative explanation for the large magnitude The negativoexcursionuas Hirstclearly seenin north-
sulfur isotope frttctionations iecorded in Ediacaran strata western Canada and southwestern United Status
that were previously attributed to disproportionation [JVarbo/i//e
e/ a/., 1994; (:arsef/fan(rKal@nan,1994]but
[ H/u ef a/., 20] 5] and is consistent with the physio]ogica] soon recognized in carbonate-rich successions worldw ide
hypothesis for the Ediacaran biomapresented here. u,ith nadar values near --8%oor lower. It is closely associ-
Complicating theview of a u1lidírectional rise in oxygen ated with tulefl rst appearance datam (FAD) of Treprfc'/rnr/.\
leveisassociatedwith the NOE is the recognition of rapid /}c'c/lr/n
[Coi'.serra
a/id Hagadol/z,2000](the 3D ichnofabric
oscillations in the õ':C compositions of carbonates presented as the pastel' child for the boundary) insofar as
(Fig. 7.2), which suggestan inherentinstability in the the initiation of widesprettdpenetratívebiotu:'bation is
carbon cycle continuing from tule SE to the end of the regarded as such a signinjcant geobiological event that it
Fortunian stage. Mtlny recent studies cave suggested tllat
is deHinedby the appearanceof such diagnostic trace
oçeansthrough this transitional interval may cavebeen 6ossiis
[/7a/zrsao
ef rz1.,2018].
stl'atified with deep ferruginous and e:)isodically euxinic h/leasurements of mdox-sensitivo traje elemento and
(containing frei H,S as in the Black Seatoday) waters the widespreadoccurrence of black share and phospho-
overlain by oxygenatedçonditions on sl)aliou' marina rites during the BACE in [ran [K//nin'a a/lí/ f]ü/a/zaóe,
pjatforms [Z,l ci a/..20 }0;Jo/?/i.\'ronef ah 2012; Spel'/hg 2001] and Onlan [Sc/nõ(/er a/fd GlorzíngeG 2007] suggest
e/ a/., 2012, 2013a, 2013b; IHood e/ a/., 2015; Bola,per widespread anoxia during this geochemicai divide, repre-
ef a/., 2017]. Fluctuations in the chenlocline could thus senting a pote11tialkill mechanism 6or the soft-bodied
dictate the spatial distribution of the Ediacaran brota Ediacaran biota, as well as their shelly rejatives C/oir(/fna
[Toirel'fr7 e/ a/., 2016], whethei' these]arge oi'ganisms were and l\ra/liacíz/af/nís [.4/}7í/tor ef a/. , 2003]. New pa]eonto-
:tnimalsthat would be aüectedby the ttbsenceof O. or logical observationsfrom southern Saberia,however,sug-
the toxicity of H:S or the osmotrophic sulílde-oxidizing gest that taxa attributed to Ediacaran and Fortunian
organisms that woujd require access to both resoulces. skeletal biomasoverlap without notable biotic turnover
Variably stratiõiedconditions may help to explain the before the BACE [Z/iz/ e/ a]. 2017J;in essence,the new
presente of "superheavy" pyrite (appioaching or data show that cloudinids and anabaritids had slightly
exceeding the ÕHScoinpositions of coevassulfato pte- longer rangemthan pnviously understood with the formei
ser\ed astraje phasesin carbonato)in Ediacaransucces- extending up through the BACE and the latter appearing
sions postdating the SE. The observation of spectacular below. ll' correct, this questíons whether the biogeochem-
34Senrichment in sedimentary sulfldes has moved some ical anomaly was a signiíicant factor in the mass
researchers to otherwise suggest that the Ediacaran extinction of the Ediacaran brota. While these shelly fos-
oceanollad low (rather than high) sulfatoçoncentrations sils aie genera1ly folded unto the broad snveepof the Ediaca ran
[Rlc'i e/ a/., 2009; Sete/lef a/. 2010; 1,OI'(/ef a/., 2012, organismo, their template-directed biomineralization

sets them apart as true animais. On the other band, the
soft-bodied forma possessed a common physiological
strittegy as desci-ibid in tais thesis: tiley may be completely
unrelated to animais. With only the possible exception
of Tira.vfa/ía and Sn'ai'//)rr/]rlír [Hagado/'n ef r]/., 2000].
thesoft-bodied forma appearnot to havemadeit across nenteuntil frei oxygencould build up in the water
the BACE.
New evidente for widespreadocean anoxia in thç pre-
lude to the BACE is interpreted from U isotope nleasure-
ments of carbonates from the Dengying Formation of
Soulh China [lr'b/ e/ a1, 2018; Z/ía/zgeí a/., 2018].]n
thesebitulllinous and fossiliferoussedimenta.Cií/ e/ a/.
[2016a] previously documented a signialcant positivo õ"C
excursionand a dralnaticshift in the õ"S of pyrite coin-
cidentwith the biological transition from Conorzrózfó-to
C/oir(/f/za. 'lltese authors suggest that an íncrease in ter-
restrial weathering fluxos of nutriente, sulfato, and alka-
linity stimulated primary productivity, biomineralization,
utndthe spread of anoxic stlbtidal and basinal environ-
ments. The new õ2;:U resulta similarly suggest the spread
of oceanic anoxia, which nlay haveplayed an important
role in the demite of the soft-bodied Ediacaran biomaand
furthei may have stimulated animal motility [Z/rr7/zg'ef a/. ,
2018]. An alternativo way of looking at the biological
arma Face that would be consistent witl] the resource-
basedhy!)othesisis that the onset of bioturbation by
mobileanimais could ha\e releasedH.S from the nlicro-
bia] mais indo the water column and there6oreallowed for
the spread of anoxia.
The driving factor(s) foi' the spread of ocean anoxia in
[he let\d-up to the Ediacztran-Cambrían bounc]ary
remains a mystery, but interrogation of the emerging Sr
isotope record of the terminal Ediacaran may shed some
llght on the problem. Recent*'SrPÓSrresults froln the fraseto the Cambrian explosion was ignited, leading to
Khatyspyt Formation in the Olenek uplift of Arctic detonation near to a positive carbon isotope anomaly
Saberiaindicate valuesaround 0.7080(Fig. 7.3), which known as the ZHUCE (Fig. 7.2) event some 1(}-15Ma
indicate a sharp declina from values that çharacterize [he
SlLand thel TES elsewhere in the woi:ld [cfu Kíizd)na/7
ef a/., 1993; Àraróa/zne e/ a/., 1994; ]Za/l,e/'sa/?er a/. , 2007].
Insofar as thesebituminous and richly fossiliferous sedi-
mentary rocks (the Ediacaran brota here are preser\-ed as
carbonaceous compressions in carbonato and sharerather
[han as casasand mo]ds in sandstone; Gla:/z(/zí/lk/n é/ a/.
2008) fere deposited at the doorstep of file Ediacaran-
Cambrian boundary, the sharp decline in s'Sr/*óSrof
ihese exceptionally well preserved limestones may record
:l significant hydrothermal event. If correct, the venting
of reduced hydrothermal fluida unto the world oceans 2014; Cbzígeo/?
ef a/., 2018]. By piercing and ventilating
may well have iesulted in the consumption of dissolved
oxygen a:ld the spread of anoxic (and perhaps euxinic)
bottom waters. Support for the end-Ediacaran rifting hy-
pothesis notably comes from Sr and Cr isotope measure-
nlenLsof TES carbonates associated with C/oz/c/friain
Unlguay[Fi'efef a/.,2011].
THEEDIACARAN-CAMBRIAN
TRANSITION123
[n sedimentary rocks deposited during the BACE,
tlranium isotopecompositions[H/e/e/ a1, 2018; Z/ia/iB'
ef a/., 20 18] notabiy reflect more oxidized çonditions, but
this may be analogousto the SE wllere ventilation of the
ocean resulted flrst in the depietion of reduced compo-
column. Molybdenum isotope measurements in basal
Cambrian sedimentary rocks in South Chintt similarly
suggest oxidizing conditions by the end of the FACE,
which could have then triggered the evolution and diver-
sification of modem animal phyla [llQ/i ef a/.. 2011]
7.2.4. The Cambrian Explosion
The global sedimentaryrecord of carbon isotope vari-
ations revealsa prolonged interval of carbon cycle insta-
bijity in carbonato-donlinated Fortunian strata, with
rapid oscillations between negative and positivo extremes
\Kttat! et «t., \99Sa\K«{\fttlal\et at.. \996\ RÍatoo4'e{ a{.,
20 ] 0], and tt riso in s7Sr/8óSrconlpositions [.Nafta/na/i e/ a/. .
1996]. It is likely that the redor landscapeof Fortunian
ocean water was similar to that suggestedfrom geochem-
ical studiesof theTES with oscillations in the chefe)online
beillg particularly important in somebasi11$
while not in
others. Analysesof sedimentary pylite and CAS in tran-
sitional Ediacaran to Cambrian sllccessions record a
signiHicant sulfur isotope sjlift [/;7ke rmd 6'l'o/zfnger. 2008;
Cz//era/. , 201 6a, 20 16c; Ha/ifó'oo ef a/. , 201 8]. Geochemic:tl
box modeting suggeststhat sulfur :enlobilization tllrough
bioturb:ttion would have enhanced oxidative processem
(inçluding sulHideoxidation) and increasedoceanic sul-
fato concentration]Car{/le/c7 a/lr7 Fargu/rar, 2009]. Wi th
[he riso of oxygenat the baseof the Fortunian. the short
latir [Küzíd)na/ier a/. 2012]
:l'hexontinuotts record of invertebiate FADA witll no
[urnover by extinction in the midst of Fortunian carbon
cycle instability is unusualand consistentwith the idea
that Fortunian ecosystems may cave been }llore tolerant
of new genetic variante [J(no// ef ír/. 1995a] and osci]-
lating redor enviromuents.This in pari may be due to the
ilovelty of bioturbation. Some of the earliest Fortunian
animais were able to penetrate the widespread microbial
mais that dominated the Ediacaran period [Àdc'/rí'oJ' a/i(/
Log«n. 1999:Bltatoiset a!., 20\4\ Cai'batia l td iNarbottne.
Ediacaran matgroundlSe//ac/ier and Py7ilgec 1994;
Boríyer ef a/., 2000], bioturbation lnay cave provided
gre:ttel' opportunities for ear:y animais by expanding hab-
itable ecosystems[À/c/7roy an(l ioga/í, 1999; Kno// a/ (r
Brr/nóízc/l,2000]. Ecosystemexpansion via bioturbation
might explain the macroevolutionarylag betweenthe
124 CHEMOSTRATIGRAPHY ACROSS MAJOR CHRONOLOGECAL BOUNDARIES

divergente of maior aniillal çlades during the Cryogetlian 7.3. CLIMATE

:utdearly Ediac:traí periods :tnd their divelsiHlcation dur-
ing the Cainbrian explosion]Ent'/n ef a/.. 2011]. InsoÊtr tts the global Cryogenian glaciations. which
Nunlerotls studies worldwide have documented an ended around 635Ma [H(!Pino/rn ef a/., 2004; Cb/z(/on
increase in the dize, diversity, and arcllitectural coin- er a/., 2005], havebeenconsideredas evolutionary bottle-
plexity of animal bui-rowsfrom the late Ediacaran to the necks [Hq/71?ian a/ü/ Sc/n'ag, 2002], it is impoitant to
eal'jy Cambrian [see G'ongeon e/ {í/., 2018 for a recent understand the temporal and spatial distribution of
compilationJ: howex-er.there is leis agreementabout the Ediacaran ice agesrelativo to that of the Ediacaran bioma,
impact of these cllanges on seafloor bioturbation. Some as well as the wild swingsin the carbon and sulfur cycles
local and global ichnological studies[l)losez' rr/zdJ?oíÜel'. that çharacterize the period. Evidente 6or icehouse çondi-
1988;Àdcliroy ati({ Log«n, \999; À.fátlgalioand Blfafois, tions through the uplxr reaches of the Ediacaran period
2014] document a stepwise increase in tule abundante aild collles fron) the preservation of diamicti tes (unsorted oi'
depth of bioturbation Ihrough the Cambrian,while poorly sorted sedimentary rocks) of conüirmed or pie-
others conclude that sediment mixing was not signiHlcant sumed glacial origin, somewith postglacial faux cap car-
until the late Cambrian [7a//ra/i a/lc/ Z)laser, 20 14] OI even bonates, aswe11asdeeply incised palco-valleys interpreted
the late Silurian [Zal'/zanef aZ. 2015]. as related to glacio-eustacy,
To assessthe geochenlicalefkcts of penetrativebiotur- Foi'emos{ alnong these is the ca. 580Ma Gaskiers
bation, high-resoiution organic carbon and pyrile sulfur diamictite in New6oulldland (a1ld its suster on the
ilnalyseswere recently conductedof the siliciclastic dom- Bonavista Península,the Trinity diamictite, which is con-
inated basal Cambriai} Global Boundary Stratotype sta:!ined to clavelasted <340,000 ye:trs; Pü e/ a/., 2016).
Section and Point (GSSP) (Chapel lsland Formation. The Gaskiers (Fig. 7.4a) is the only Ediacarandiamictite
Newfoundland, Cartada). Along these wave-waslled to cave thus far beendirectly dated by U-Pb techniques.
exposuiesat Fortune Head (giving a narre to the basal Notably, the short duration of the Gaskiersice age is
Cambrian stage), a positivo õ"C exculsion in organic inconsistent with ])redictívedurations for snowball Earth
tllatter was noted to start at e.racf/ the Ediac:tran- evento, altllough a low-latitude (19.1 t l l.I') paleom:tg-
Canlbrian boundary alld returns to stably "C-depleted netic determination [P/sal'('t-skJ' ef a/., 201 t] for the
values hundreds of metera higher coincident with enric1l- Avaloll tear:tin in whicll it sinsand a thin yet discontin-
ments of t)C in carbonato carbon [oward seawater va]ues uous cap carbonato (see inset) with a negativo-to-positivo
lf7a/ir.toaeí a/-, 20] 81.Pyrite in tllese sedimentanotably õi3C trenó [iW]'p'oit-a/id Kai{/nzan, 1999] are consistellt
underwent signiflcant "S depletion at the Ediacalan- with other aspecto of Cryogenian global glaciation.
Cambrian bouildary that continued upsection towt\rd the Many researcllerscave attenli)ted to correlate the
end of the Fortunian. This observation is suggestiveof Gitskiel-s giaciatlon with the SE as recorded in middle
!he growth of the oceanic sulfate reservoir spurred by Ediacaran carbonatesworldwide [see review in X7ao
progressiveventilation and oxygenation of shallow sedi- et a/., 2016], but the potentia} 6ar repetition [Acfc/üdr/en
ments as a direct consequente of penetrative bioturba- e/ a/., 2008] or diachroneity of the biogeoçhemical
tion, which further stimulated the oxidative sulfur çycle. anomaly and the laçk of post-Cryogenian glacial diamic-
These authors suggest that sediment ventilation in the tites in critical successionshave severely hindered our
basal Fortunian spun'ed a tel)lpoi'ary inçreasbin MSR abilityto demonstrably connect çlinlatiçand geoçhemieal
and benthiç sulfur çycling under low-oxygen conditions. evento. Based on their lithological characteristics and
A declina in bottoin water pO, could claveresuited from indirect radiometric constrítints, othel potencial Gaskiers
illore efllçient biological ptlmping, and MSR play have equivalente include the Mortensnes [Ha/ erson e/ a/.,
generated the organic carbon isotope excursion that ter- 2005] and Moelv diamictites [2?frzgen ef a/.. 2005] in
nlinatesin the upper Foitunian strata. The end of the northern and soutllern Norway; theHankalchough [XI(in
organic carbon isotope excursion implies stabilization of ef a/., 2004]:\nd Hongtiegot}[S/ie/i e/ a/., 2010]diamiç-
c:trbon and oxygen cycling in the shaÍlow substrate, tites in northwestern China; the Egan diamictitein
potentially related to a balancebetween the production, Australia [Co/'ke/'on a/l(/ G'eorge, 200 1; Carkeí'on, 2007]
export. and remineralization of oiganic matter and/or to and the Croles Hall diamictite in nearby Tasmania [Ca/!,el'
a stabilization of valer column !)O.. Overall, these data e/ í//., 20041;the SerraAzul diamictite in Brazil [H /lrrle/iga
from a siliçiclastic dominated successionshed new light e/ a/., 20071;and tlte Squantum diamictite in the Boston
on the Cambrian explosion. They attest to the geochem- Basin, United Status[T7zo/n/}.fo/l a/i(/ Bota'r/ng,2000]
ical signiHlcance of the initiation of sediment ventilaiion Ediacaran sucçessionsin theseareítsare typically domi-
by animais at the dawn of the Phanerozoic,with oxygen- nated by siliçiclastics [//CZ#lnanan(/ Z.i, 2009], which com-
;ition of the shallow substrato soon followed by {lte plicate their conelation with carbon cycle ttnomalies
tippearance of biomineralízed small sheljy fossils (SSFs). typically recorded in carbonato Ihcies. Nonetheless,
 (a)

Figure 7.4 (a) White to red colored Gaskiers diamictite overlain by a 50cm thick white carbonato (left of the field
lssistant)on the chore oí Conception Bayat Harbour Main, Newfounclland. Insetshowsthe brecciated a1ldpotentially
karstified upper surfaceof tulethín carbonate filled with green rlludstone of the overlying Drook Formation.(1))Green
meta-basaltof the basal Catoctin ft)rmation with diapirs (flame structures)of faucluier formation carbonate injected
behveei] the chilled pillow margenson Goose Creek cear Aldie, Virginia, Uníted States.Inset illustrates a hyaloclastic
texture at contact between the Catoctln meta-basaltand marble of the fauquier cap carbonato, indicatlng that the
sedimenta were water-saturateclduring emplacement of the volcaníc rocks. Reproduced u,ittl permíssion of Elsevier.
(c) A 20m thlck diamictite along the Tas-Yuryakh River lylng unconformably above Turkut Formation dolomites in the
Olenek uf)liR, Arctic Siberia, Russia.'he freshly exposed dÊamictite has a green-gray sandy to clayey calcareous matrix
with abundant cobble- to boulder-sized clasts in a weakly stratifíed pide.The randomly oríented clasts(see inset)are
primarily derived from the Turkut ar)d un(Jerlying Kllatyspyt formations, but they algo collsist of occasional green
igneous rocks and metarllorphic rocks of exotlc origin. Subrounded clasts plucke(f from the surrounding
carboilate-rica matrix are notam)lyíaceted.(See/nserffo/' co/or /epresentaf;onof fhe f7gt/re.)
126 CHEMOSTRATIGRAPHY ACROSS MAJOR CHRONOLOGICAL BOUNDARIES

strongly to moderately negative õ"C compositions al'e f\4 ountains of southern Siberia C/oiro/íia hax e been found
noted in postglacial carbonates above the H:utkalchougl}, in diamictite nlatrix) and :tre inteibedded or lie belos
Egan, and Serra Azul diamictites, and thesehaxe been those containing elemento of the basal Fortunian
provisionally correlated to the SE. Paleomagnetic data A} abctritestais\llcalttsZoxne.
notably support a low-latitude position fol' both the Egan Supporting the view of a boundary ice age,a 20 m
t\nd Hanki1lchough diamicti tes [/;r(Z/#}7í a/zd Z,i, 2009], thick diamiçtite [Kat{/}llan eí a/., 2009]composedof stlb-
wllich, assuming synchronicity of the presumedGaskiers rounded to angular cobble- to boulder-sized clasts (many
equivalents, would blue the lides beta'een Cryogenian and faceted) of underlying lithologies in a gray-green sandy
middle Ediacaran ice ages. to ciayey calcareous matriz was recently iecognized along
Radiometric, stratigraphic, and paleolltological data the Tas-Yuryakh Reverin the Olenek uplift of Arctic
from Laurentia, Africa, China, and Saberiafurther sug- Saberia (Fig. 7.'k). The Tas-Yuryakh (!iamictite was
gest the possibility of post-Gaskiers Ediacaran ice ages, deposited unconformably above carbonates of the Turkut
:tlthough there is scal'ceevidente 6or striated ciasts, till Formation (similarly containingelemen
ts of the .4. /i'üz//-
pellets. or dtopstones in most of these deposita. For c-afn- Zolle) at the sandelevei where a volcanic tufo was
exanlple,in northern Virginia, United States,dianlictite identiHiedand radiometricallydated at 543t l IWa
of the Fauquier Formation is stratigraphicallyoverlain [Bott'l'üig ef a/. 1993]. Tltis weakly stratiHiedpile includes
by a thiçk red bed s:tndstone ttnd a 20 nl tlliçk carbonato seven disLinct leveis interpreted tts lodgnlent till [HI/zai/d
horizon that preservesa strong negative-to-positivo õ"C and EJ;/ex,2002;Hi/iarr(/,2008] that acculntllatedat the
twnd [/7ê/)er/ e/ a/., 2010].The carbonato htts a con6orm- interface of a grounded ice sheet in a shi1llow marina
able soft-sediment contact with piliow basalto of the environment. Lacking evidente tbr tectonic uplift and
overlying Catoctin Formation (Fig. 7.4b), which is con- erosion of the plat6orm. it is likely that incision of {ulder-
strained by a U-Pb zircon age of tlround 571Ma. lying li thologiesresulted from glacio-eustaticlou,ering of
Application of the Catoctin age to the Fauquier diamic- sea levei by 150m or more. 'Hie paleokarst is a prominent
lite supports the vier of a regiollal glaciationminemil- lliatal surface that shows systematic variations in the
lion years younger than the Gaskiers [seealgo l,/17ne/lrc//l/i anlotult of erosion throughout the Olenek uplift
ef a1, 201 8 for a bloadly equivalent post-Gaskiers glacial [Pokroi'.vkJ' rr/i(/ h/iag/'adot', 1991; Pakr -skJ' an(/
diamictite in pera-Gondwanan West Africa] and within Nlissarzltevsky, \9q3\ K+toiiet cii.. \99Sb\ Petechatyet a!.
uncertainty of tlte earliest fossiliferous Ediacaran strata 1996] with progressivodowncutting of the Turkut and
in Avztlonia. An even younger Ediacaran ice age (the equivitlent strata across tule btisin. Tais uncollformity
Vingerbreek glaciation) is suggested by the presente of partittlly er:\$es the piofound negativo õt3C excursion that
deep a1ld laterally extensive palco-vajleys interpleted to characterizes the Ediacaran-Cambrian boundai'y
Lave 6o:'med during glaçio-eustacy and Hiiled with a het- (Fig. 7.2) in tuleOlenekuplift and is expressedthioughout
erogeneous mix of !imestone-clast congiomemte, gray- the eastern Siberian platform [K7ro/}?enfav.\-kl'.
19901.sug-
waçke, quartzite, share, carbonate, and tillite (with gesting glacio-eustacyas zt potencial driver. However, a
abundant faceted pebbles). Radiometriç constraillts diatreme that cuts tlirough ullderlying strata is recog-
[Grp/z/reger'
ef a/.. 1995] supporta ca. 547Ma agefal' the nized along the Khorbusuonka and Mattaia rivers, which
is the likely sourceof the dated volcanic tuas, and
Vingerbreek event in Natnibia and its potentia! correlativo
in Souto Africa [Sc/ni,e//rnrsí1
9411Ge/:/nsi1972f1983; undoubtedly contributed to th(pglacially remobilized
Gei't}s an(! Grcsse, \99\ \ PI'aekctt et {tt., 2üQ8\Get'tusalt({ volcano-sedimentary sucçession. The Tas-Yuryakh
Gaiíc-#er,2012]. If the drawdown surfãce is correctly tied diamictite is immediately overlain by mixed carbonate
[o the regional buiidup of ice sheets (as consistent with a and siliciclastic rocks of the early Cambrian Mattaia
moderate negativo shift in carbonate õ''C values at the Formation, which preservemtrajes of animal activity as
baseof the member; Say/nl e/ ír/.. 1998), the Ediacal'an wel! as diagnostic SSFsand reef-building aj-cileocyathids
brota wotild have felt its chiliing ef6ects. representing the most ra])id phase of animal diversifica-
Fina1ly, glacial deposits or deep paleo-valleys believed tion in Earth's }listory [Rozano\,ef aZ, 1969]
to be related to glacio-eustacy neitr the Ediacaran- The observationsof a widespread Baykonurian ice age
Cambrian boundary cave been described fro m thio ughou t suggest a glacial divide between the enigmatic Ediacaran
central Afia [see review in C7izí/nakot',2009], West Africa biomaand the succeedingCambrian explosion. Insofar as
[Bei'rlarz(/.Sar:/:z/f ef a/. , 1995], tutd Namibitt [Gein7s. 1972, oxygen was tt criticamresource for the Ediacaran biotit.
1995]. These ice age deposita are best known from cordel TES temperaturas associated xx'ithicehouse condi-
Kazakhstan aild Kyrgyzstan and are collectively pari of tions (starting with the Gaskiers e\ent and continuing
the Baykonurian glaciation. Sotlle of these glacial episodically toward tule Fortunian) would cave further
deposita occur above sedilnentary rocks containing saturated seawaterwith the brealhing gas.While increased
Ediacaran fossils (in one case from tlle East Sayan O. concentrations would be consistent witl} the "com
 THEEDEACARAN-CAMBRIAN
TRANSITION 127

cradle" hypothesis [ }lcke/i-R/c/E. 2007] for the brota, tule cl[ finges, tl]e evolution of the Ediacaran brota is a!] origins
breathinggas is 28x leis solublethan CO, and 83x lesa stop'y based on the ride of oxygen in seitwater. On one
solublethan H.S at 20'C. which would havellad a hand, the oxygencould cavepromoted eukaryotic metab-
signiHicanteüect on the pH and toxicity (to animals} of olisllls including the forillation of sterols [R!//7negar.
the valer column and pare fluida. Notably, a report of 199]; Car/f/zge/ aZ. 2005;Bu(/d,2008] and of collagen
Mo isotope variations across tule Ediacaian-Cambrian [Tolt't', 1981; Sair/, 2009], but on the other the ride of O
boundary in íman and South China [ll''f//e ef a/., 2008] could have stinlulated autotrophic sulhde oxidation that
suggeststhat H,S play cave spread across shailoxx'deposi- led to the evolution and diversification of mat-like oigan-
tional environments,venceproviding a potencialkill isnls connected to a common metabolic strategy.
mechanism for the Ediacartul brota, assuming these My initial interest in the preservation of Ra?lgeír,
organismo fere animais. Alternatively, the coup de grâce /'fel'/(//r?/lí/ll, a/i(/ Eni/e//a stemmed from the observation
may cave been simply the drawdoxx'n of oxygen as ocean of yellow coatings coillposed of jarosite, a pi'oduct of the
:inoxia spread in the TES [see X.ülnf/a anc/ 14/a/a/iate, oxidative weathering of early díagenetic pyrite [l)a/'/-ac/l
2001; Z/ia/ig ef a/, 2018], especiallyif the biota was ef a/, 2012; Ha// ef aZ. 2013], on freshly exposed fossilif-
dependenton sulHideoxidation as a common metabolic erous surfaces (Fig. 7.5a; seealgo Fig 7.6d as presented in
strategy. Recognition of a boundary glaciation plovides a colo! in ric'kers-Rfc/r ef a/. , 201 3). Astrobiologiçal interest
physical mechanism for sea-levei drawdoxx'n and the injarosite stems from its presence on the surface of Maré.
enhanced upwelling of H,S as well as ''C-depleted HCO.' tx,hich suggests wet, acid, and sulfato-rica conditions
prod uced through sulfato reduction in deep anoxic waters, early in planetztry history [Sqzíyl'esef rz/., 2004]. ]n
which contributed to the strong negativo õ':C excursion association with theseEdiacaran fossils,the p:esenceof
preserved in marina carbonates. In turn, the ecospace jarosite is arguably related [o the grota'th of pyrite "death
vacatedby the Ediacaran brota may cave been repopu- masks" [Ge/1///7g,
1999; Hn(/t?/-xon
ef a/., 20 11], w]lich he]p
[ated with Canlbrian faunos in the g]acia] aftermath, in LO explain their tmusual preservation as exquisitely
likely responsoto rising oxygen leveis in the ocean aild detailed imp:'essions in í'ine sande and silos [but see
ltmosphem. /Ver:/?ra/í e/ a/. , 20] 6; 7?n/ran er a/., 20 16 for an alternative
explanationfor mat and fóssil preservationbasedon
7.4. ERNIETTAVILLE:EARLY BIOTURBATION? silic:t saturation of the Ediacaran oçeiln)
While the deatl] mask hypothesisis particularly attrac-
The resouKe-based hypothesis for the ride and fal! of tive, there is no ít priori iettson that the living OI'gania
tule Ediacaran brota was 6arnlulated in the midst of che- tnembranesfere not at leastpartially pyritized while the
mostratigrapllic and paleontological research of lower oiganisnls were alive. Given how rapidly the organic
Nama Group (Kuibis Subgroup) sedimentary rocks in sheets would decay, there must have been available H,S
southern Namibia just east of Aus on Farm Aar an(] the and Fe:' in poi-elluids to supportmineralizationin bife.
surrounding region. Observatioils of the shallow m:trine (3iven the remarkably low solubility of both hematite
Nama Assemblage in southe!'n Africa were augmented (X',. -10's) and pyrite (.K.. -10''''), an even partially
by severalexcursions to Mistaken I'oint in Newfoundland oxidized water colunln would have l)eenlacking in these
to study the deepwater llabitats of the lona-ranging reduced aqueous species. This vier is consistent with Fe
Avalon Assemblageíuid to South Australia to vier and speciation constraints tilat suggestthe Ediacaritn bioma
describe the cosmopolitan White Seu Assemblage. lived below w:\terátltat fere at le:tstepisodically oxic and
In all but onedebatablebasin [À/ac(/oníz/r/
ef at. 2013: low in irotl [lHoo(r ef {z1., 2015]. Tlte sedimenta be]ow the
seediscussion ttbove], the Ediacaran brota appear in the ubiquitotls mais, however, would necessarily have been
stratigj'aphic record after the SE, which on several cotlti- anoxic in ordem
to supportanaerobicMSR andits output
nents presei'ves cotipled chtt ages in file carbon, sulfur of hydrogen sulHtde.Under reduçing conditions, Fe'* pre-
and strontium isotope composition of carbonates[C#f served as oxida coatings on detrital nlinerals could cave
e/ a/., 2015 and referentes therein] that together suggest been reducedand mobiiized within poro fluida, which
[he riso of oxidants in seawater and of oxygen in the could then reaGEwith hydrogen sulHlde to form pyrite
:ttmosphere. These changes fere most likely driven by on the fractal interior walls of the oig:tnism. Interior
Pan-Afriçan orogeny and silicato weathering that deliv- pyritization could caveconferredan advalltageto the
ered sedimenta, nutrients. and oxidants unto the oceítns. Ediactuan brota by providing structural support for
promoting primary productivity and organic carbon the organismo or to regulatethe fme diflusion of O. to the
burial IK {Pnan e/ íz/.. 1993; Sqrrire ef rr/., 2006; Oc'/ra/?d inferior of the organism.With a predominantly endoben-
S/r/e/(7s-Z/íozi,
2012; P/anal'xA.v,2018], as well as the ride tllic iifesty:elCi'flziei a/zd/t(/onA-úí,1996; Gr rz/r(Zz//ik/na/i(/
of su!!'ate concentrations [Fike e/ a/., 2006; Kazfánlan Se//íz(#zúE 2002; Sef/ac/rer a/z{/ G'fs/l/fck, 2014], it is the buried
ef a/., 2007]. !n light of these coupled biogeocllemical portion of the organismsthat would be initially pyritized
Figure 7.5 (a) Freshly exposecl Pterfd/nlu/n surfaces coatecl with yellow colored jarosite, a hyclrous sulfate of
potassium and iron that forma as a product of pyrite weatherlng, at Aarhauser on Farm Aar. near Aus, Namibla
IHa// et a/. 201 31. 1nsetshows a typical iron oxidize patina on an exposed arlcl weathered surface of Pferidfniurn
at the somelocalít},.Reproduced
with permissionof Elsevfer.(b) "l\4u(fchip" l)recciaassociatedwith Ernleffa-
bearing sandstone from Frnietta Hill on FarmA.lr. The mud chips cave the iron oxida p.atina indicated above, arld
a partíally exposed erniettid is exposed on the surface (see red arrow). Thís suggeststhat the chips rcf)resent the
surface connection between iníaunal Ernletfabasesand theír epibenthic fronds. (c) Sock-shapedsanclstoneconcre
bons with flat rappersurfaceslikely to representErnfertaspecimensthat have lost thcir tubular covering lhrough
exposure and wenlhering on \Vinda Punk, Farm Aar.These specimens are the some size and general shape to Ern;efta
l)reserved in situ with tul)Miar structures (d) (yellow scale bar l cm). (See InsereÁorco/or represenrarlorr oflhe /lgt/re.J
 THE EDIACARAN-CAMBRIAN TRANSITION 129

b.

.:J-

&

'q'''iT'

Figure 7.6 Exceptionally preserved Erra/erraand Rangia specimens cliscovered in sandy gutter castaon FarmAar,
southern Namibia. (a)'üe most complete frnietta specimen known to date [/t,antsovet a/« 20161, wíth white
arrow pointlng to the positíon of the connectíon behveen infaunal l)ase anelepibenthic frond (see Fig. 7.5b: com
is 2.26cm in diameter). Reproduced with l)ermission of Wiley. (b) lllustr'ated reconstruction of the frn/erra
specimendepicted in (a) with tubular l)ilayer shown. Reproducedwith })ermissionof PeterTrusler.(c) lllustrated
reconstruction of /?angeawith a tul)ular core and sixfolcl symmetry of vares inclu(Jingat leastthree ordensof
fractal folcling. Reproduced with permission of PeterTrusler. (d) The most complete Rarlge l specimen known to
date IVTckers-Ríc/7 ef a/., 201 31 showing san(f-fil led baseof the organism (inlage of some specimen in Vickers-R/ch
ef a/. j201 31 shows patina of yellow jarosite, a weathering product of pirite)
130 CHEMOSTRATIGRAPHY ACROSS MAJOR CHPONOLOGICAL BOUNDARIES

Other saclike, bulbous. baggy, and pimpled iloldfasts [ubes that form a palisac]e-like structure, which 6orm a
illighl then refleçt e:irly pyritization and the requirement zigz:tg sutule at the base of the band-filled anchor.
of the mttt-like OI'ganiam to miga't\te within tule sediment Extending beyond the bag-like base,the par:\llel tubes
in arder to harvest additional sulflde.Tule migration of continue unto two hcing faniike structures (Fig. 7.6a ttnd b)
the organic walls would be fal' moj'e }ealistic if the The connection of the two parta (see wllite al'row in
interior' (rttther than the exterior) of the organism were Fig. 7.6a) would representthe position of the mud chips
preferelltially pyritized. Pyi'itization in life couid discussed above. Other representativosof the Nama
explain why ET'/r/e/fabasesare se common and \À'llythe Assemb[age, inciuding Pari'í(/ffz/ir/}l and ]Va/;ia//a, are
Ediacaran fóssil record, in general, is rei)lote witll hold- strikingly similar to E//i/t //a being composedof pali-
fasts but piopoitionaily few'er fronds. Supporting this sades of tubos, but in larger and more complex boas and
argument, dose examination of specimens pleserving bela shapes]Graz/r(Zz/nk//!
aní7 Sef/ac/it'r, 2002]. In çon-
bota holdfast and ftond reveals torsion in the latter. t!'ast, Ra/igeaconsistof an axial bulb and stalk that
wllile the foi'mer appears cemented in plane [Perel' extends indo a cyjindrical cone; the axial strt:ature lk)rms
TI.zí /er, pera. Coinm., 2018; see Fig. 7.7e). Whlle l)lany the foundation for six vales arrangedradialiy around the
of the Ediacaran taxztc]ear]y ]ived above the sea bot- axis. with eachvane consistingof z\ bilaminar sheetcom-
ton} and were buried and preservedduring storm posed of repetitivo tubular eienlents exilibiting at least
evento [ }/7c#el's-Rfc/i ef a/.. 201 3; /pa/i/sol, ef a/., 2016]. three ordensof self-similar bianching (Fig. 7.6c and d)
[hey nonethelesshad ho]dfasts that connected trem to The common tubular constructional elementoof the
the substrato, allowing for a direct connection to the Ni\ma Assenlblage, including Su.arrpif/iria [iVa/'bo/ine
H.S resource witllin anel beneath the tllats. er a/. . 1997], suggestan adaptation to an infaunal lifestyle
Evidente 6or the pyrite molda, however, \ç'eathers in order to maximizegtts exchangeand nutrition
quickly away onde fossils are exposed, ol'ten leaving [GI'í/z/i(/a/ikfrr an(/ Seí/ac/ler, 2002, 2005]. In this senso, the
behind il'on-rica surfaceveneels (Fig. 7.5a inset). In the growth of tubular bioHilms from the sedimenta (siphoning
case of Ei'nlt f/a. the veneer is usually t)lissing, leaving microbially producedH.S untothe fronds whereO, would
bellind bulbous, lunlpy, and rock-shapedsandstoneballs. diíTusein from the \x'atercolumn through osillotrophy)
eachwith a single round to elliptical tlat surface.These provides a mechanisnl to gather the two critical resources
specimens (Fig. 7.5c), which !itter tlle glounds wllere neçessary to sustain tl)ioautoti'ophy, teus poteiltíttlly rep-
well-preservedisolated and in situ specimensof tlle same resenting the earliest form of bioturbation (conrl-ízZ):/k,
dize and shalw, but with tubular external inorphology 1999). The double layel-sof parallel tubosin Ei/?/erraand
(Fig. 7.5d), :tl-efottnd, are interpretedto iepresent the its relatives nlight play physiological foles with one deliv-
interior sandstone Hills of the organic-walled organisms. e!'ing H,S to the fi-ondaand the other O, to the holdfast.
Recal]that thin walls and inert interiors would aliou the One of the fundamental properties of biological mem-
surface arett/volume of tule Ediacaran biomasto approach brai)esis tllat theyare barriersto the pernleationof
[hat of osmotrophic baçteria. Furthermore, Hieldsof sub- polar moleculesresultingfrom the fact that the paraf-
lllat sandstone surfaces associated with individual Hinic interior of bilayer lipid meinbranes is hydrophobic
Ei /zfef/aspecimens are similarly llkely to be the preserved [ }y/(/o/nsAíre/ a/., 2007]. A]though H,S is s]ight]y pcl]ar
expression of coiltinuous undulating sheets of organic and has somellydrogen-bonding
capability,it too is
menabranesrwith the top surfaces typiHied by ovoidlild llydrophobieíwhieh lendato its toxicity'as it will readily
oxidized "lllud chip" breccias (Fig. 7.5b). Given tracei of partition indo cells [R/a/r/ a/ií/ Rou'/cy, 2014]. The perme-
tubular impressionaon the surfaceof the chips ttnd tule abitity of H.S throughbiologicalmembranes is tour
co-occurrence of partiam Ernfeffa specimens emanating ordens of magnitude mole thítn water but is less than
from them (see ar!'ow), the lnud chips al'e atternatively that of nonpolar O, [Su/urzynsk/ef cz/.,1989]by 0.5 to 2
interpreted as the surface expressionof connections bet- ordens of magnitude based on experimental [A/ar/ra/
ween individual E/'/7/er/«bases and their fronds or of er a/. 2014] and lnodel c:\lcuiations [Rfa/i/ a/?(/Ran'/Q',
fields of fi'onda emanating from a single expansivo sheet 2014]. The bottom linfa is that if the thin walls of the
of under-mat organic structures. Ediacaran fronds were similar to lipid membianes, they
On Farm A:tr, completespecimensof EI/ifef/a and vx,oulddiRusein O, faster than the H,S cotlld escapeotlt
Rangeu XÀ'eretransported during storms and exquisitely although the rales woulcl undotlbtedly be clependenton
presewed in sandy channels and gutter casta [HÍ('kei'i- the concentration gradiente i1lsideand outside of the
Rf(/í e/ aZ, 2013; E//fo// e/ a/., 2016;/I'a/r/ o},e/ a/., 2016]. organisms and their elTective surface áreas [Z,(Ü7a/pr/íe
The discovery of these unique lagerstãtte allowed for the e/ í//., 2009]
most detailed reconstruction of file organismo to dttte The irregularities of nlicrobial mata reílected in the
( Fig. 7.6). .Er/r/cf/a has a saç-shapedbo(ty with w:\lls çon- varied expiession of microbia1ly induced sectimentary
structed of two pa:allel layers of vertically arranged structures (M ISS) [/VQ#ke ef a/.. 200 1] suggest the potentia]
(a) k'3u.N (b)

Figure 7.7(a) Conception-sty]e preservation [Narbonne, 2005] of Charnfodfscus spinosus aria Cyc/)ruç procerus in
positive relief on a surface of the Mistaken fbint Formation of Newfoundland. The fossíl surface is overlain by a thin
bed of volcanic ash (vinil)le asdark layer on the upper right). (b) Spindle shaped Fracfo/usas177israi
with primary and
secondary branches on the somesurface in Newfoundland with visible volcanic ash. Fossilsin (a) anel(1))are mem-
bers of tuleAvalon Asseillblage. Imagem(c-f) are from the Ediacaran member in South Australia and archived at the
South Austral ía Museum (while scale bars are 5 cm in length). 'Íhese are representativos of the White SeaAsseml)laje.
(c) Dlckinson/a cos(ata moving (see arrows) anel resting trajes. (d) Kfml)ere//a quadrada (rlght arrow) anel assoclatecl
trace fóssil Kfmberfch/}us feruzzl {left arrow). (e)Áspfde//a wlth holdfast anelstalk, but no frond.(f) ,'\rborea arbolea.
Source.'Government of South Austraiia. (Seeinse/f áorcalor represa/ faffon of f/TeÉi'gare.J
132 CHEMOSTRATÉGRAPHY
ACROSS MAJOR CHPONOLOGICAL BOUNDARIES

Ihat tuhes could fol'n} through wrinkling itnd then propa- and fractally constructedorgailisms flourished in
gate i1lto sediments. While microbial mata are noi-mally oligotrophic and bacterially dominated marine environ-
composed of horizontally strtltiHled communities, they nlents. The most parsimollious interpretation then would
calaexhibit sedimentary structures deílned by dynaillic be that the Ediacaran biota themselves were either bacte-
physioctlemicalgradiente,as well as the diversity t\nd rial colonies or, more likely givei} their complexity and
physio[ogy of its denizens [Kri 7be//r ef a/.. 1994]. inulticellularity, that they symbiotically hosted sulflde-
Oxygenic photoautotiophs usually occupy the tlighest oxidizing bacteiia within their bodies [cf:, Bzrrzyrnk/
levei in the mais, while anoxygenic photoautotrophs and É/ a/., 20171.
chemolithoautotrophs lie belleath, and these microbes N4oie problematic are the Ediacaran oi-ganismsthitt
utilize H.S as an electron donos in the reduction of ttppear untethered to the substrate, have been çharacter-
carbon dioxide to organic manter. Of particular concern ized as bilaterian,and/or are associatedwith feeding
hereaie the non-p1lotosynthetic sulfide oxidizers that use tracei, including dickinsonids, sprigginids, parvancori-
tllolecular oxygen from tulew:\ter column and sulfide pro- nids, :tnd the monospeciHic genus of Kf/z b( rc'//a [Gt/i/f/zg
duced lower in the mat through MSR [Z/le/c-:fr7.çkaia ef a/., 20 i4; DI'a.çel'a/lí/ G'e/l/frlg,20 í 5]. Notwithstanding
e/ at, 2014] to sustain their nletabolic activities.While ttrguments suggesting that specimens lacking bilateral
photosyntheticsulfide oxidizers are also possible,any symmetry are taphonomiç variante, norte of theseforms
common metabolic strategy 6or the Ediacaran brota have a demonstrable mouth, gut, or ânus. The nlovemen{
would have to accommodaie the deepwaterforms living tracei of l)/ck/n.\o/i#r(Fig. 7.7c)and its relativesllave
far below the photiç zune, especiallyrepresentativos
of been interpreted asevideltçe for osmotrophiç feeding on
Ihe Avalon Assemblage (Fig. 7.7a and b). [lle mata [],aÜ/an]/ne e/ a/., 2009], but these O]ganisms
According {o Fi'/eí/r'/c/r ef íz/. [2001] and referentes might alternatively move between fE'esh surfaces in an
therein, file biological oxidation of llydrogen sulflde to eüort to quickly absolb H,S unto their systemsin arder to
sull'ateby a phylogenetically diversoarray of prokitryotes rechal'ee the resourcefor the sulüideoxidizers within.
is one of the major reactions of the global sulfur cycle. Given tlle permeability of the sour gas througl] biological
Sulfur oxidation in [he Architea is restricted [o some melnbranes. this processmight bettel' explain the preser-
while bacterial vation of bota movementand Festinstrajes.On the other
tnembersof the arder S!/!/b/oba/e.ç,
oxidation of sulfur is mediatedby both anaerobic plloto- hand, Kfrnbere//a[seeFig. 7.7d] has beenintei'pieted asa
Irophs aild aerobic lithotrophs. The latter is tulemetabolic mollusk:tn-grade organism [.fk(/o/ik/n írn(/ H/aggoner.
strategy considered here givei} the likelihood of a terminal 1997] and is doctlmented in dose association with fans af
Neopioterozoic buildup of oxygen in the {itntosphere and scratches(seettnow) interpreted as trt\cesof nlat grazing
oceans,t\lthough other oxidtults are known to play a role [Fc'(/o/lkfner a/.. 2007]. Hera it is the absenceof an ânus
in this nletabolisin [G/roó'/z
rr/ld Díí/n, 2009]. Aerobic and/or of Ediacaran
coprolitesthat posesa problem,
sulfur-oxidizing bactéria belong to neai'ly 15 known which could be solvedif the proposedsiphon tllat
venera, including .4c'f(/fr/r/obac///us, Beggfrz/orz. and scratched the mat was designed to evacuate it of H,S
Tllese autotrophic bacteria Hix carbon rather than organic nlatter. It is difHicultto slloehorn
P.çerí(h/planas.
dioxide to crente 6ood either via the reductive pentose these complex and apparently motile forma with those c)f
phosphatecycle or via the reductive tricarboxylic acid sessile infaunal and fractal grades [Z)/'oóc'p'a/lc/ Ge/í/frzg,
cyele-NVith the ability to hariless energy and produce 20151:but tlleir bilateral symmetry is debatableas are
[heir own 6oodthrough su]üideoxidation, the ear]iestsed- issues related to their physiology, if there was a common
iment miners arguably developeda wide vztriety of bau- inetabolisill sha!'edby all of the Ediacai'an brota, then the
plans to accommodate environmental gradiente. For unoi thodox speculations entertained heracould explain tt
exame)le, the /'«ngeo/Pior/u/r.ffrom Avalonia might cave lot about their lllorphological disparity (i.e. regou!'ce
had to signinlcantly increase their fractal surface :u'e:ts gathering in diH'erent sedimentary and basinal regimes)
and/or their height above the sediment-waterintei'face as well as their demite.
(Fig. 7.7a and b) in arder to deal with lower O: abun- l t should not be overlooked that sulíide-oxidizing sym-
dantes in deep bottom watt:'s [A]'f//se/ a/., 2014]. If bionts are harboredwithin rare modem aninlals living
correct, eçoiogical tiering of the Avalonian bife forms under unusuitlçircumstances[Nt-/ó'o/ia/ic//Tx#er, 19951
would be a strategy for ilarvesting oxygen at diüerent For example, tubo wonlas like R#?/a /ua(/iD,/////a (from the
leveis above the substrate rather than organic molecules polychaete fitmily Si/)og//rif(/ae)living Real'hydrothermal
ICtaphal} af:d Nal'bolltle, 2QQ2;Ghisalbct'íi cf a}., 2Q\ 41.In vents [Bp'fg/if a/?(/ l.a//f( z', 20 10], gutless phallodriline oli-
support for the donlinance of pl'okaryotic s111Hide- gochaete xx'aulasliving in oxygen-denicient I'egions of the
oxidizing communities in the TES, biomarker studies inocean [B/azq/akef a/.. 2005},and frei-líving nenlatodes
Báltica indicate unusually lügh hopane/éter:tnerácios within the family Dc'ó'nio(/or/dae
[Be/X/n e/ a/. 2018] all
[Pc/rr e/ a1. 20 18]. This nnlding suggests that these large beneHit from symbioses with sulllde-oxidizing microbes.
 THEEDIACARAN-CAMBRIAN
TPANSITION 133

While the symbionts within the tule worms at deep ocettn geochemicttl, ecological, and physiological in6ol'mation
ridgesutilize H,S emitnatingfrom black snlokersand O. In their 2013 review, Laflítmme and colleagues conclude
directly from seawater,thosein marginal l arine settings that behavioral innovations associated with predatioi]
beneHit fronl the ability of the host animais to migrítte and ecosystem-wide ch:tnges, reflected in the matground
between sulflde- atld oxygen-rich sedimentary environ- to nlixground transition (Fig. 7.1), likely spelled the death
tllents. Other modem animais (e.g., À c'le/x) {tj-e known to knell for the Ediacaranbrota. In this cttse.tlte Hirstvertical
tolerate high sulHide enviromllents, and these might be perforations of the matgrounds near the end of the TES
important modem analogs for true Ediacaran meta- would ha\e dramatically increased the flux of ieduced
zoans. It lias been suggested that st1lfur-based nletabo- gasesto seawater,including both H,S and CH,, tllat
iismshelped chapeinicial symbiotic evento,leading to the would have combined with available O. and hí\stened the
evolution of both unicellular and multicellular eukary- spiead of deepoceananoxia. Evidente for anoxia around
o\es tOI'et'niífnn and van Getnetefen,20(n; Tlieisseii et af. tule boundary comes fioill traje element, as we11as Mo
2003; /t(fe/i/( / a/i(/ luar//n, 2008]. itnd U isotope studies [K/pmr/'a an(f PHíífa/urbe, 2001;
Scln'õdcr «nd Gi'ot=íllgci',2001; !yiife et a1., 2Q08; ttQI
7.5. METAZOANS
TAKETHESTAGE t-f a/., 2018; Z/gange/ r//.. 2018 and referencestherein],
although the incompleto preservation of the stratigraphic
While bioillarker and possible fossil evidente suggestsa t-ecord in bota South China and Oman complicates the
pre-Ediacaran origin of sponges [Lol'e e/ a/., 2009; timing of evento as recorded in widely separated basins.
À/a/o(y'ef a/., 2010; Bloc-ksef a/- 20i6], Ediacaranevi- [f su]fide oxidation was theçommon metabo]icstrategy,
dente for the earliest veriHiablemetazoans comes in the [he declina of oxygen at the boundary wou]d have wiped
6orm of Doushantuo embryos preserving a range of out the Ediacaran brota, at the sande time promoting
cleavagestages, diH'erentiated multiceljularity, and the MSR and the j)roduction of authigenic carbonato(either
;tbsence of a çell xx'ê1ll.as well as {he presente of diapause in the wttter cohlmn or in sedimenta) strongly depleted in
eggcysts [.otan e/ «/., 1998, 2014; yi/í ef a/., 2007]. Tule ';C [Sc/n'age/ a/., 2013; Crrí ef í//., 2017],which coutd
:animalalTinity of these fossils has nonethelessbeen chal- explain the tlegative Ediacai-an-Cambrian boundary õ"C
lenged by observations along the present-dayNamibian excursion noted in most sucçessionsworidwide
coast of gitlnt sulfide-oxidizing bactéria of the menus
r7z/a/na/'grrp'f/íz]Baf/ey e/ a/., 2007], which are quite 7.6. CONCLUSIONS
unlikely to be fossilized[Cn/lnfng/la/}z
e/ a/., 2012]. Tule
sudden appearitnce of template-directed biomineraliza- In Older to explain the stratigraphic and spatial distri-
tion in the cloudillids, whiçh appeai'worldwide in car- bution of the Ediacaran biotas, one nlust plane trem in
bonate facies some time afte! the SE, may be further thc enviromnental context in which they fere found.
evidente for the earliest Ediacaran metazoans. Otherwise. From a chemostratigraphic point of view, !he largo and
clear evidente of animal activity comes from wormlike fi-actally constructed olgiinisms, post of which fere
organismothat crawled througll the mais using peristaltic rooted in the substratoamong the ubiquitous microbial
nlotions that left increasingíycolnplex trace fóssil behind mitts, :tre tightly flamed between two of tlte most pro-
ICa/'Z)onea/ií/ N«róo/ine, 2014; A/eye/' e/ a/., 2014; 6ound negativo çarbon cyçle anomalias in Earth's history:
ScJlj[ybaifaelas..2Q1 6; Bttddatld Jcnsen,2Q
\ ]]. ]\\ese Lhe older SB and a younger event neta' the Edi:!caran-
ilnimals apparently inined the mata for nutriente and O., Cambrian boundary known as the FACE. Assulning
but they would also cave to have contencled with abun- these bookend geocllemicaleve:lts had enviromnental
dant H.S if they ventured too deep[C7le/?
e/ a/., 2013]. In drivers and are not simply globally distributed diagenetic
tais case, these early metazoans would cave to be at least artif.acts, the spatial and temporal distribution of the
episodicítlly tolerant of the toxic gas, unless they too har- Ediaçaran organismoshould be understood in termo of
bored sulnide-oxidizing cheillosymbionts. changes in global ocean chemistry, enlerging ecological
The cii'amaticend of the Ediacaranbrota near the opporhlnities, alld evolutionary innovations.
Ediacaran-Cambrian boundary may represent the Hn'st Researchers often seek out specific morphological char-
gre:tt illítss extinction, the biotic replacement of soft- açteristics of the Ediaçariul brota in order to fi{ these enig-
bodied metazoans by Cambrian aninlals, or tlleir gradual matic organismo unto prefelred interpretations as animal,
disappearance in the fóssil record as a result of the elimi- plant, lichen, or microbial clades, with Adolf Seilacher sug-
nation of the matgrounds in which they thrived [Zad/ünz/lzr gesting that the brota Hjtsunto a phylum distinct froi)l any
ef aZ.. 2013]. The latter is counteled by the pl'eservation of modem representative. Given the similttrity of primary
Ediacaran-style mais and enviromllents in basal tubular and fl-actallyHoldedunits in the constructionof
Fortunian strata [Bu«ro/se/ a/., 20141,but evtt]uating the t1leseorganismo, one nlight çonsider whether they {t11had a
other possibilities requins an approach that integrates unifying metabolic strategyto explain their morphological
134 CHEMosTRAriGRAPHY ACROss MAJOR CHRONOLOGiCAL
compjexity rather than trying to Hlt strangemorphologies list of card-carrying paleontologists
Isto recognizable categorias. 30+ years have entertained my questiona and listened to
The resource-basedllypothesis entertained hera stetns my often heretical ideal about the life acrossthe Ediacalan-
from geochemical blues for both the riso of O, in the oceano Cambrian tmnsition. Foiemost amotlg theseare Andy
atld atmosphere in middle Ediacaran time and the
continued production of H,S in the sedimenta.Expanding
upon the vendobiont hypothesiswith the likelihood of
multiceilularity and symbiosi$ tule geochemical blues as
well as the morphological constructlon tuld disparity of
lhe Ediacaran biota aie consistentwith the utilization of
these two critical resources t'equired for thioautotrophy.
These organismo are envisioned to have absorbed oxygen
from the water column and pumped hydrogen su16idefrom
Ihe sediments in order to sustain their unique physiology
:uld explain their nlorphological peculiarities. If trufa.the
Ediacaran biomasvele anlong tlte earliest ecosystem engi-
neers.Their search for resources both in the sedimenta and
in the water column would represa:ltthe earliest 6orln of
bioturbation. which would cave propagateda shift in the
chemical and ])hysical properties of the sedimenta
[Sr/rãg/)mrcl'
e/ a/., 2016]. Furthermore,insofar as their
occupation of the subsurface environillen t delivered auto-
trophiç organic mattel- to the sedimenta, theil activities
would have connected the pelagic to benthic iealms Horthe
Hlrsttime in Eart1l'shistory [ET'n'üíe/ a/., 2011].
Whether a heterotrophic animal, eukaryotic symbiont,
or bizarre bacterial bife 6orm is dependent on sulHKle
oxidation to make a living, the declina of oxygenand
spreadof hydrogen sulflde untothe water column cear to
the Ediacaian-Cambrian boundary would spell trouble
6or any organism that could not move away from toxic
conditions [C/re/i ef a/. 2018; Z/ra/zg ef íz/. 2018]. With
the assumption that these bizarro lide forma were not ani-
maIS the biotic replacement hypothesis would not apply,
leaving mass extinction of the Ediacarail experiment the
post i[ke[y exp]anation for the bio]ogic:t] crisesat the
Ediacaran-Cambrian boundary. Given my predilection
nocgeocllemisti.y, the spread of:anoxia :!nd loas of the O,
resourcecear the end of the Ediacaian period remam my
preferred interpretation; however, there could be ttn
iilternative (or suppjementary) cause for the mass
extinction, assunling thesetubular' and fi'actaspneunlatic
oiganisms fere unusual bacterial coloilies or complex
eukaryotic baús that housedsulfide-oxidizing bactéria. In
either case,predation by the eillerging army of mcltile and
carnivorous metazoanswould cavepunctured the organic
walls of the Ediacaran brota, thereby deflating the H,S-
üilledba11oonsand leading to their ultinlate demite.
ACKNOWLEDGM
ENTS
The author wishesto thank the editols and publisllers Bailey, J.V.,Joye,S.B., Kalanetra, K.M., Flood, B.E., and
of tais iinportant volume,especiallyAlciciesSial and Ritu
Base, tor their insight and patience, as well as to a launclry
BOUNDARiES
who ater the past
Knoll, Gerard Gernls, Guy Narbonne, David Bati;jer.
Fr:\nk Corsetti. Steve Rowland, Shuhai Xiao, Zhou
Chuanming, Nick Butterfield, Dama Grazhdankin,
Patrícia Vickers-Rich. Tom Rica. Peter Trusler. Les
Kriesfeld, DONSErwin, Maoyan Zhu, Jim Gehling. James
SchifR)auer, Gregory Retaliack, Tom Holtz. John Merck,
Simon Datroch, and Mare Laflamme. The ideas enter-
tained in this chapter are my own and are not intended to
throw shade on any of my esteemed colleagues who no
doubt could enectively cotmter many of my argumenta.
Thanks Risoto the three external reviewersand to a wide
range of internationa{ Hieldgeologists, stratigraphers, geo-
chronologist$ and paleomagnetists u-ith whom l haxe
w,orked (and argued with) over the decades as we Êocuson
[lle peculiarities of the Ediacaran and Cambrian world.
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