The Ediacarian Period and System: Metazoa Inherit the Earth

Author(s): Preston Cloud and Martin F. Glaessner

Source: Science, New Series, Vol. 217, No. 4562 (Aug. 27, 1982), pp. 783-792
Published by: American Association for the Advancement of Science
Stable URL: http://www.jstor.org/stable/1689058
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 27 August 1982, Volume 217, Number 4562 SCIENCE

Semantic impediments have made un-

ambiguous discussion difficult. Until
1930 no term existed for all of Earth
history characterized by metazoan fau-
nas and evolution, while pre-Cambrian
The Ediacarian Period and Syste: mi: was the only inclusive term available for
the long preceding interval which, here-
Metazoa Inherit the Eai rth tofore, had yielded no convincing rec-
ords of visible animal life. Chadwick (9)
then proposed a solution: the geological
Preston Cloud and Martin F. Glaes ssner record characterized by conspicuous an-
imal life would become the Phanerozoic
(Greek for visible plus animal life),
whereas antecedent history and rocks
Ever since Darwin, the geologically presence in these rocks ( of glauconite and would be called Cryptozoic.
abrupt appearance and rapid diversifica-other sedimentary pec uliarities indica- Phanerozoic is now widely accepted,
tion of early animal life have fascinatedtive of a marine origi in bolstered the but Cryptozoic has had only limited use.
biologists and students of Earth history expectation that, if thei re were anteced- By definition, the term Phanerozoic
alike. Yet, until recently, convention and ents to the Cambrian bi otas, they would must be extended downward to include
lack of data limited serious discussion of be found here. older discoveries of manifest animal life.
planetary evolution to its last eighth, That prognosis was cc )nfirmed with the In the opinion of one of us (P.C.) this
with emphasis on details of metazoan discovery by Sprigg in 1947 (1) of what should take the Paleozoic with it, adding
and tracheophyte evolution, sedimentol- has come to be called t :he Ediacara fau- a basal extension to previous additions at
ogy, and paleogeography, and with but na, together with its description by the younger end of the original Paleozoic
passing reference to earlier Earth his- Glaessner and others (2- -5), and with the Era. Chadwick's definition, geologic
tory. recognition that Ediaca] ra fossils are old- consistency, and etymological congruity
all equate the base of the Phanerozoic
Eon with that of the Paleozoic Era. Inas-
Summary. The Ediacarian, here defined as the initial period arnd system of the much as the initial Phanerozoic rocks
Phanerozoic Eon, is characterized by the oldest known multicellula ir animal life. The and history are also pre-Cambrian, still
distinctive biotal assemblage comprises naked Metazoa, represe,nted in the type older divisions of rocks and history can
region by 26 species in 18 genera and 4 or more phyla, plus simple netazoan
r surface no longer be unambiguously designated
tracks. Elements of this unique biota appeared worldwide at lo' w paleolatitudes, as simply pre-Cambrian. Clarity de-
following terminal Proterozoic glaciation. Ediacarian history lasted I from about 670 mands that the term Cryptozoic be ac-
million to 550 million years ago. This interval, plus Early Cambri; an, was the time cepted or pre-Phanerozoic employed
la and most of the where the intent is to designate by one
during which metazoan life diversified into nearly all of the major phyl
invertebrate classes and orders subsequently known. word the long sequence of rocks and
history that preceded the appearance of
metazoan body fossils, imprints, and
This constraint loosened with the er than and different fi rom those of the tracks as conspicuous components of the
growing perception that some seven- conventional Cambrian l (4, 6). Further geologic record of life. The other writer
eighths of geologic time had already support came with the d
liscovery of simi- (M.F.G.) considers it unlikely that the
elapsed before the Cambrian Period be- lar faunas in ancient roc :ks of southwest- great majority of geologists will refrain
gan. Increased knowledge, both of the ern Africa (7) and new finds in the En- from using the entrenched formal term
extent of time and of the older rocks, glish Midlands (8). It i s the gist of this Precambrian.
stimulated reconsideration of the nature article that this Ediaca tra fauna charac- Discussion relevant to the base of the
of biological processes during those long terizes a distinctive sode
epi, of pre-Cam- Cambrian (and, by implication, the Pa-
eons that preceded the Cambrian and brian history but of Lnerozoic
Pha age and leozoic) became active from the late
accelerated the search for evidence con- that, among names ava Lilable, it is most 1940's through the 1960's (10-12). It
cerning them. Well-preserved sequences appropriately called theEdiacarian Peri- grew in scale and scope with research
of sedimentary rocks, at places extend- od. Rocks formed with iin this period of and discussion within the framework of
ing far beneath the Cambrian without history then constitute\ the Ediacarian
apparent major interruption, showed System, and the appear; ance of this fan The authors are, respectively, Senior Queens Fel-
ance of this fauna low at the Baas Becking Geobiological Laboratory,
ansition from the Canberra City, ACT 2601, Australia, and Professor
that there was neither a great historical marks the geological tri
Emeritus, Department of Geology, University of
discontinuity nor a universally high de- preceding Proterozoic ,on
E to the follow- Adelaide, Adelaide, SA 5001, Australia. Dr. Cloud's
gree of metamorphism beneath the Cam-ing Phanerozoic Eon, comprising the current address is Department of Geological Sci-
ences, University of California, Santa Barbara
brian, as once supposed. The common rest of geologic time. 93106.

SCIENCE, VOL. 218, 27 AUGUST 1982 0036-8075/82/0827-0783$01.00/0 Copyright ? 1982 AAAS 783

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the International Geological Correlation ous strata of the Wilpena Group. The
Program (IGCP), established by the In- distinctive fauna is one of soft-bodied
ternational Union of Geological Sciences arthropods (Praecambridium and Par-
and UNESCO in 1971 (13-16). As a vancorina), segmented worms (Sprig-
result, the conventional base of the Cam- gina and Dickinsonia, Fig. 2, C and D),
brian has moved stepwise downward primitive colonial cnidarian coelenter-
from being defined by the first appear- ates (Charniodiscus, Fig. 2A; Glaessner-
ance of the trilobite Olenellus to nearly ina, Pteridinium, and Phyllozoon), a pos-
coinciding with the massive incoming of sible proechinoderm (Tribrachidium,
shelly fauna-mainly small primitive Fig. 2E), and some dozen genera of early
mollusks and tubular to conical fossils of medusoids (Fig. 2B) and other simple
uncertain systematic position. coelenterates. It is widely distributed
A broad consensus has also emerged through an interval that reaches a thick-
concerning the grounds on which useful ness of 112 meters near the middle of the
historical-stratigraphic subdivisions are Pound Subgroup at Mayo Gorge 15 km
best made (11, 14, 17). The agreed upon north of Hawker (19) (Fig. 1). The fossil-
criteria are biological, with emphasis on iferous portion of the Pound has been
the ranges in time of marine invertebrate called the Ediacara Member (20).
animals where dealing with the Phanero- On fossil evidence from the type se-
zoic. More attention has focused on quence the base of the Ediacarian is at
boundaries than on the modal character- least as low in the Wilpena Group as the
istics or distinctive contents of the divi- middle of its upper clastic division, the
sions bounded, perhaps because the re- Pound Subgroup. A new discovery of
gional variations are so great. Whether Fig. 1. Index map for key localities in Flinders simple metazoan tracks in the Elkera
Ranges. Stippling indicates outcrop of Pound
for boundaries or for modal characteriza- Formation of north central Australia im-
Subgroup. Stars mark Ediacarian fossil local-
tion, however, it is clear that paleobotan- ities. [Adapted from Wade (19) and Jenkins plies a still lower position-at or beneath
ical, paleomicrobiological, and other cri-and Gehling (48)] the base of Bunyeroo Formation (21).
teria must also become important as we Combining paleontology with paleocli-
undertake to define the lower limits of matology, operational convenience, and
Phanerozoic and Paleozoic rocks and In brief, the Ediacarian Period and the probability that the metazoan record
history and to delineate older divisions System-named from and characterized will be extended downward to some ex-
of the historical and sedimentary record. by fossil evidence now known from tent, we find it more appropriate, at this
Granted that the purpose of nomencla- some dozen South Australian localities time, to equate the base of the Ediacar-
ture is to facilitate unambiguous discus- along almost continuously exposed lines ian with the base of the first stratigraphic
sion, we must somehow specify the se- of outcrop 140-kilometers long (Fig. 1), unit above the highest tillite-containing
mantic content of terms utilized and seek and with equivalent strata worldwide-is deposits beneath known Ediacarian fos-
to eliminate or clarify words that now not synonymous either with the officiallysils. In South Australia that is the con-
mean different things to different people.defined Vendian of the Soviet Union or formable lower surface of the Nucca-
the Vend of general usage. The Ediacar- leena Dolomite and equivalent rocks at
ian System is the oldest unit definable bythe base of the local Wilpena Group,
Ediacarian Period and System metazoan fossil content. In its stratotype overlying the tillitic Elatina Formation
area it follows conformably above tillite- of the Umberatana Group. From data
In this spirit, we turn to the matter of containing late Proterozoic sequences, now available, this could be of roughly
how best to treat the historical episode but it is succeeded disconformably or the same age as the terminal surfaces
represented by the Ediacara fauna and unconformably by fossiliferous Lower of other ice-related deposits that oc-
its equivalents. It turns out that this Cambrian deposits. cur shortly below faunas of Ediacarian
fauna not only has a circumglobal distri-We then define the Ediacarian Period aspect in other parts of the world (22,
bution, but it is clearly antecedent to as that interval of geologic history char- 23).
conventional Cambrian. In fact, the term acterized by the soft-bodied, macroscop- Biological support for placing the
Ediacarian has already been applied to ic, marine invertebrates of the Ediacara boundary so low is the structure called
this part of Earth history-as the Edia- fauna and allied forms. Rocks of the Bunyerichnus (Fig. 2F), found near the
carian Stage of Termier and Termier (12) Ediacarian System range from immedi- middle of the Brachina Formation some
and the Ediacarian Period of Cloud (16). ately above the last preceding episode of 1800 m below the main Ediacarian fossil
In the present article we seek more clear-glacial sedimentation in South Australia zone (3). Although we agree with Jenkins
ly to define and document the historical and, with important exceptions, the last (24) that this surface marking is not a
episode, to consider its appropriate posi- of the fossil stromatolite Conophyton true crawling track, we do not agree that
tion in the geologic time scale, and to beneath equivalents elsewhere, to the it is "unlikely to be of metazoan origin"
review alternative nomenclatures. Other massive onset of shelly fauna and dis- (25).
names that have included some or all of tinctive trace fossils (for instance, Sko- Thus the basal Ediacarian approxi-
the rocks and history we here call Edia- lithos, Rusophycus, Plagiogmus, and mately coincides with the oldest record
carian either have not been clearly de- Phycodes pedum) that marks the initial of manifest animal life. For those who
fined, have meant something different Cambrian at many places. may still harbor reservations about that
from the type Ediacarian, or have fluctu- In the stratotype area of the Ediacar- conclusion, it should be noted that most
ated too much in meaning to be useful ian System, described below, these (P.C. thinks all) so far reported pre-
(18). rocks are represented by the fossilifer- Ediacarian Metazoa are pseudofossils,
784 SCIENCE, VOL. 217

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are misdated, or are not Metazoa. Some Type Site and Global Reference Section subdivisions are in the middle third of
are even modern organisms or their the Parachilna sheet between Mayo
traces intruded into older rocks (16, 26). The discovery site for the Ediacara Gorge and Brachina Gorge, along the
Among recently proposed examples, the fauna is at the southern end of the Edia- western flank of the central Flinders
structure called Rugoinfractus, suggest- cara [also Idracourza, Idyakra, or Eti- Ranges (Fig. 1). The general stratigra-
ed as evidence for a metazoan presence kaura (29a)] Hills near Randell Lookout,phy, paleogeography, and correlation of
1100 million years ago, appears to be the southwest corner of the 1/250,000 Cop- the rocks of the Wilpena Group and
fillings of shrinkage cracks in underlyingley map sheet (- 30?50'S, 138?8'E). related strata in this and adjacent areas
strata (27), a common form of pseudofos-Most Ediacarian fossils so far described are summarized by Preiss et al. (30).
sil. Other supposed ancient Metazoa from Australia have come from this area, Descriptions of the individual units ap-
cited by Durham (27) from the Huronian but only a small part of the sequence of pear in papers by others (19, 31).
Ajibik Quartzite of Michigan and the interest is exposed there. We designate the steeply northwest
early Proterozoic Medicine Peak Quartz- The thickest fossiliferous sequence, dipping (55? to 60?) sequence in Bun-
ite of Wyoming are also of inorganic 112 m, is in Mayo Gorge at the southwestyeroo Gorge (- 31?25'S, 138?32'-35'E),
origin, as at least one of their authors has corner of the 1/250,000 Parachilna map 380 km north of Adelaide, as the stan-
now recognized (27a). sheet, whereas the type sections for the dard reference section for the Ediacarian
Concerning prior usage, Termier and Wilpena Group itself and all its named System (Fig. 3) (32), with the sections in
Termier (12) proposed "L'Ediacarien,
premier 6tage paleontologique . . . car-
acterise par ces fossiles et qui s'insere au
sein du sous-systeme Eocambrien." De-
spite their assignment of Ediacarian tc
Eocambrian and some erroneous corre-
lations (for instance, with the much older
Belt Supergroup of Montana), there is no
doubt that the Termiers intended the
Ediacarian to comprise the initial Pha-
nerozoic strata, characterized paleonto-
logically. Here, moreover, although we
follow Cloud (16) in elevating this term
from stage to period rank, we also com-
ply with standard Phanerozoic nomen-
clature in adopting Ediacarian for both
geohistory (period) and rocks (system).
Although biostratigraphically defined, it
is not a biostratigraphic unit (a zone) in
terms of the current International Strati-
graphic Guide (28). In those terms it is a
chronostratigraphic (system) and geo-
chronologic (period) unit.
Division of the Ediacarian into formal
epochs of history and series of rocks is
not proposed here, although it may be
suggested by faunal variation. Cribri-
cyathids of Early Cambrian aspect
(Cloudina) occur with Ediacarian forms
through much of the lower Nama Group
in the southwest Africa, whereas typical
Early Cambrian trace fossils (such as
Phycodes pedum) mark the overlying
Nomtsas Formation and the Fish River
Subgroup (29). By contrast, strata of the
Wilpena Group have so far revealed nei-
ther shelly fauna nor such advanced
trace fossils. This, with its unconform-
able base, suggests (to P.C.) that the
Kuibis fauna of the lower Nama Group
may be younger than the Ediacara fauna
proper and that, if this should be sub-
stantiated, subdivision of Ediacarian on
faunal grounds into epochs and series
may become possible. Until then, the Fig. 2. Some distinctive Ediacarian Metazoa (A) to (E) are from Ediacara Hills. (A)
Charniodiscus arboreus (Glaessner), plaster mold, photography by R. J. F. Jenkins; (B)
divisions shown as Early (Lower) and
Cyclomedusa radiata Sprigg; (C) Spriggina; (D) Dickinsonia costata Sprigg; (E) Tribrachidium
Late (Upper) Ediacarian on our global heraldicum Glaessner. (F) Bunyerichnus dalgarnoi Glaessner, from near middle of Brachina
correlation chart are provisional. Formation in Bunyeroo Gorge; compare with (B).
27 AUGUST 1982 785

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the Brachina Gorge and Mayo Gorge as Geochronology was glauconite, which may be detrital or
supplementary reference sections. We formed by alteration of old biotite, giving
choose Bunyeroo Gorge as the strato- The available geochronologic data for exaggerated ages for enclosing sedi-
type (32) because (i) an uninterrupted rocks of the Ediacarian System have ments. Such a number, for example, led
sequence of some 3000 m of Ediacarian limited usefulness. Some papers in to the erroneous report of a Proterozoic
strata is here compactly exposed in al- which ages are cited do not state the age for the Cambrian trace fossil Sko-
most continuous outcrop within a hori- method employed, the kind of rock or lithos in the Wessel Group of Northern
mineral dated, or the decay constant
zontal distance of less than 5 km, (ii) the Australia (33). Finally, international
usual fossiliferous interval is well dis- utilized. Where such data are available, agreement on decay constants was only
played, and (iii) the oldest known record the method used was often potassium- reached in 1976. Before then potassium-
of likely metazoan life (Bunyerichnus) argon dating, which is susceptible to argon, rubidium-strontium, and even
occurs in this sequence. The Brachina error in both directions. Argon loss withuranium-lead ages included a computa-
Gorge section is more accessible by car, time and metamorphism results in mini-tional variance among laboratories of
and the incomplete Mayo Gorge section mal ages for materials dated. Other K-Ar several percent.
adds information on the stratigraphic numbers may be too old because of Cowie and Cribb (34) sought to ame-
range of the most distinctive Ediacarian absorption of argon by thermally altered liorate such problems for the Cambrian
fossils. pyroxenes or because the mineral dated by choosing 40 selected radiometric ages
and recalculating them to the new decay
constants. These were plotted on a cali-
brated chart to ascertain average num-
bers for Cambrian, Ediacarian, and relat-
ed rock boundaries. The number so ob-
tained for the base of the Cambrian was
560 million years if defined as the top of
the Tommotian Stage, or 590 million
years if placed at its base, a placing that
we favor. We have reservations, howev-
er, about the validity of dating statistics
that depend so heavily on K-Ar ages of
glauconite.
A more credible age for the Ediacar-
ian-Cambrian transition comes from as-
sessment of recent data for the English
Midlands: Nuneaton, Warwickshire;
Charnwood Forest, Leicestershire; and
the Wrekin area of Shropshire.
The Nuneaton ages are on distinctive
diorites ("markfieldites") that intrude
(and are thus younger than) the Calde-
cote Volcanics of supposed latest pre-
Cambrian age. At Cliffe Hill in Charn-
wood Forest these "southern diorites"
also intrude strata containing an Ediacar-
ian type of medusoid fossil (35). This
locality provided most of the samples
from which a Rb-Sr isochron age of
552 + 58 million years was obtained for
the southern diorites by Cribb (36), re-
calculated to 540 ? 57 million years with
the 1976 decay constant. That is close to
the 546 ? 22 million years found for sim-
ilar South Leicestershire diorites some
10 km distant. In the Nuneaton area the
Caldecote Volcanics are unconformably
overlain by the Hartshill Formation,
containing fossils of the Baltic Lower
Cambrian Mobergella zone 250 m above
the volcanics (37). These Lower Cambri-
an fossils are thus younger than the age
of 540 million to 546 million years found
for the diorites that intrude rocks with
Fig. 3. Aerial view of stratotype area. Abbreviations: -Ch, Cambrian, Hawker Group; -Chp,
Parachilna Formation (basal Hawker Group); E, Ediacarian System; Ewpr, Ediacarian, Ediacarian fossils.
Wilpena Group, Pound Subgroup, Rawnsley Quartzite; Ewpb, Pound Subgroup, Bonney
In the Wrekin area, about 100 km west
Sandstone; Eww, Wonoka Formation; Ewbu, Bunyeroo Formation; Ewa, ABC Range Quartz-
ite; Ewb, Brachina Formation; Ewn, Nuccaleena Dolomite; U, Umberatana Group (Proterozo- of Charnwood Forest, Patchett et al. (38)
ic). North is at the top. [Courtesy of the Department of Lands of South Australia] obtained a good Rb-Sr whole-rock iso-
786 SCIENCE, VOL. 217

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chron age of 533 + 13 million years on
the Ercall granophyre, unconformably
overlain by fossiliferous Lower Cambri-
an of uppermost Tommotian to Atdaban-
ian age.
The numbers and stratigraphic rela-
tions in these three Midlands areas thus
reveal an episode of igneous activity
around 533 million to 546 million years
ago, following deposition of most of the
Charnian strata of Ediacarian age. Much
but not all of the Lower Cambrian, how-
ever, is younger than that. In order to
Table 1. Taxonomic affiliations of metazoan tive Ediacarian elements are present at
fossils from the Ediacarian of South Austra-
this place, but ages and local succession
lia. Numbers of additional taxa from central
strongly imply an Ediacarian equiva-
Australia are added in parentheses. Percent-
ages refer to specimens collected at Ediacara. lence, and imprints of soft-bodied worm-
like metazoans occur in these sediments.
Phylum
The numbers given above are believed
andhclass Genera Species
and class
to be the best so far available for Edia-
Cnidaria (67 percent) carian rocks. They imply a range in time
Hydrozoa, 3 3 from perhaps 670 million years at the
Chondrophorina
base to around 550 million years at the
Scyphozoa 4 (+2) 4 (+2)
Conulata 1 1 top.
Others (medusoids) 3 (+ 1) 6 (+ 1)
Colonial Cnidaria 4 5

find a realistic age for the base of the Annelida (25 percent) Metazoa
Polychaeta 3 7
Cambrian we must add some years to Arthropoda (5 percent) 2 2
these numbers to compensate for miss- Phylum uncertain 1 1 The distinctive fauna of the Pound
ing Tommotian sediments. Estimating (3 percent) Quartzite at Ediacara (Fig. 2 and Table
that missing time as 10 million to 20 21 29 1) has already been described (1-5, 48)
Trace fossils 6 7
million years and averaging, we find 550 and tabulated (49). It consists entirely of
million to 560 million years as a likely 27 36 remains and imprints of soft-bodied Me-
age for basal Tommotian and hence the tazoa and trace fossils. Some show indi-
Ediacarian-Cambrian boundary. cations of chitinous or minor spicular
That number is not far from earlier strengthening elements such as are found
accepted estimates of around 570 million of nearby Finmark in similar in livingnorthern
soft-bodied inverte- Norw
years. It differs significantly from the a number close to those reported for brates, but none had mineralized shells
estimate of 590 million years given by Soviet diamictites of presumed Varan- or solid skeletons. About two-thirds of
Cowie and Cribb (34), for basal Tommo- gerian age and glacial origin (42). Ages the specimens known are cnidarian coe-
tian, based mainly on K-Ar ages for around 650 million years are commonly lenterates. Such a dominance is rare at
glauconite from the Soviet Union. These suggested for the Varangerian else- younger pre-Cenozoic fossiliferous lo-
ages, however, are increasingly being where. Thus it is possible either that calities and unknown in the Cenozoic.
questioned by Soviet authors. these mainly K-Ar ages are too young Unique though they be, however, fossils
Concerning numbers relevant to the (because of Ar loss), that they date meta- of the Ediacarian assemblage are clearly
base of the Ediacarian, Coates and Preiss morphism rather than sedimentation, or related to younger fossil and even living
(39) reviewed Rb-Sr data for the ages of that the north European tillites are in factforms.
glacial and overlying sedimentary rocks younger than the ones that give older Rb- Hydrozoa are represented by the dis-
in Western Australia, converting them to Sr dates in Australia. In any event, the tinctive floats of the Chondrophorina.
the new decay constant X87Rb = 1.42 top of the terminal Proterozoic tillites Among them is Eoporpita, belonging to
x 10-11 year-'. They concluded that cannot at present be dated more closely the same family as the living Porpita,
three shales of lower to middle Ediacar- than between - 670 million and 650 mil- colonies of differentiated polyps that
ian age above the uppermost glacials lion years. drift at the surface of the sea. The only
gave reasonably good Rb-Sr whole rock In Newfoundland, Hughes and Bruck- other living genus, Velella, resembles
isochrons. Two of the three, correlated ner (43) suggested that the intrusive Ho- the Ediacarian genera Ovatoscutum and
with the lower Brachina Formation of lyrood Granite, with an adjusted Rb-Sr Chondroplon, although these have bilat-
the stratotype Ediacarian, give ages of age of 594 million years (44), is also of erally symmetrical floats. A number of
672 ? 70 million and 670 ? 84 million about the same age as the Harbour Main other genera of similar colonial hydrozo-
years, respectively, compared with a volcanics and the Conception Group of ans occur in the Paleozoic, but only two
less well constrained date of 676 + 204 marine volcanogenic sediments. The lat- still live. It seems that either competition
million years on the Brachina itself. A ter contains in its upper part the Mistak- among planktotrophic Metazoa or preda-
younger shale equivalent to the Bun- en Point fauna of Ediacarian affinity (45), tion on them increased with time.
yeroo Formation midway of the strato- found well above tillites but below the The Scyphozoa are apparently repre-
type sequence gives an age of 639 ? 47 Random Formation and equivalent or sented by such genera as Brachina,
million years. older rocks with Tommotian fossils (46). Ediacaria, Rugoconites, and Kimber-
Other dates of interest include those Similarities between the Mistaken Point ella, considered to be a possible ancestor
related to fossils of Ediacarian aspect in fauna and the Ediacarian of Charnwood of the Cubomedusae. The Conulata, an
Scandinavia, Newfoundland, and North Forest combine with lithological resem- important group of Paleozoic fossils, are
Carolina. Kulling (40) illustrated simpleblances to imply equivalence and pre- represented by Conomedusites, striking-
trace fossils and a medusoid (Kullingia drift proximity. ly like the Ordovician Conchopeltis. A
concentrica Glaessner) from beds corre- The firmest number available for rocks number of medusoid fossils, particularly
lated with strata above the uppermost that probably represent part of the Edia- the common Cyclomedusa and Medusin-
Varangerian tillites and below Early carian System is a U-Pb concordia age of ites, are not sufficiently distinctive to be
Cambrian fossils (Volborthella) in north-620 ? 20 million years on zircons from placed in classes and orders of the Cni-
east Sweden. The age is younger than anlittle metamorphosed pyroclastic rocks daria. There is, however, little doubt that
adjusted Rb-Sr isochron age of 654 mil- in conformable stratigraphic sequence most of them lived like common modern
lion years reported (41) for the Nyborg with fossiliferous volcanogenic sedi- jellyfish.
Formation below the uppermost tillites ments in North Carolina (47). No distinc-Sessile colonial Cnidaria have left
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 The wide distribution of Ediacarian
faunas and even species supports paleo-
geographic reconstructions that place
fossiliferous localities of the Proterozoic-
Phanerozoic transition interval in low
latitudes (54). Equatorial currents fa-
vored by such reconstructions would
have facilitated the observed spread of
marine faunas.
The Proterozoic to Early Cambrian
sedimentary succession has been inten-
sively studied in recent years in search of
data on which a stratigraphic definition
of the base of the Cambrian might be
based. It is clear from this work (15) that
the Ediacarian fauna of soft-bodied ani-
mals disappeared from or became very
scarce in the preserved record after
Ediacarian time. In its place appeared a
fauna of small shelly fossils, many diffi-
cult to place in the zoological system of
Fig. 4. Distinctive microbial fossils. (A) to (D) Micrhystridium sp. from lower Yudoma Group
(Ediacarian), right bank Aldan River, upstream from Belaya (or Xanda) River, East Siberia. (E)
classification (5). They are not descend-
and (F) Obruchevella parva Reitlinger from Muraykhah Formation, Jubaylah Group (Ediacar- ed from members of the Ediacarian as-
ian or Cambrian), Jabal Umm al'Aisah, northeastern Saudi Arabia (59). semblage. Their ancestors may have
been too small to be fossilized or may
not have developed mineralized tissues.
strikingly leaf-like fossils, up to 1 m long. The status of the Ediacarian as a valid That this post-Ediacarian fauna with
Some of them (Charniodiscus, Fig. 2A)Phanerozoic chronostratigraphic unit of abundant and diverse small shelly fossils
show distinctive characters of the living system rank in the standard global strati- should be considered Cambrian, despite
Pennatulacea (Anthozoa, Octocorallia), graphic scale is reflected in this fauna its lack of trilobites, is supported by the
while others (Pteridinium, Phyllozoon), and reinforced by its position in the occurrence in Siberia of small, primitive
although pennatulacean in some re- geochronologic scale. It is characterized archaeocyathids at the base of the Tom-
spects, are more difficult to interpret. by the global distribution of distinctive motian Stage. They soon developed
New specimens of Rangea (50) have metazoan assemblages resembling those complex calcareous skeletons which
shown it to be an endemic Namibian found at Ediacara and differing from built reef-like structures in post-Ediacar-
genus to which the first finds of this those of the succeeding Cambrian Sys- ian time. Their evolution through the
group of fossils from Ediacara were mis- tem. The recognition of their potentiality Early Cambrian to their extinction in
takenly assigned. The term "Petalona- for long-range correlation began with the early Middle Cambrian time illustrates
mae," proposed by Pflug (50) for these discovery of Charniidae and medusoids the successes of minor groups during the
and other Namibian leaf- or cup-shaped comparable with those from Ediacara in initial Phanerozoic adaptive radiation of
fossils and for similar Ediacarian genera, the Charnian of the English Midlands (2- the emerging Metazoa.
has proved confusing and unnecessary. 5, 8, 48, 49) and related forms in south- Like other transitions in the geologic
The significance of these fossils is the western Africa (7, 29). Charnian types of record, however, the replacement of
wide geographic distribution and the fossils were later found, together with Ediacarian soft-bodied by younger skele-
great age of ancient colonial cnidarians, others, in the Mistaken Point fauna of tal Cambrian faunas was neither com-
some of which closely resemble living Newfoundland (45) and in Siberia (52). A plete nor geologically instantaneous. The
seapens (for instance, Pennatula), orga- remarkable display of this fauna is that inevolution of chitinous cuticles and of
nisms that are rare or missing in most of the Valdai sediments of the White Sea tubes formed from discrete grains ce-
the Phanerozoic fossil record. Coast of the Soviet Union (53). It in- mented with mucus started during Edia-
About 25 percent of the specimens cludes about ten species and ten genera carian times, and we note reports of
collected at Ediacara are annelids. The (not all represented by previously known Sabelliditidae with organic tubes, possi-
most common genus, Dickinsonia, may species) also found at Ediacara. A simi- bly representing Pogonophora, of puta-
have survived into Paleozoic time (51). A lar assemblage occurs on the southwest- tive Riphean to Early Cambrian age. The
ern margin of the East European plat-
similar form, Spinther, is still living as an characteristically Lower Cambrian crib-
ectoparasite on sponges. Spriggina is a form. The Soviet faunas include numer- ricyathids are represented by the genus
very different kind of free-living, proba- ous Dickinsonia costata, Tribrachidium, Cloudina in calcareous strata of the
bly nectobenthic, polychaete worm. and the primitive arthropod Vendia, sim- Ediacarian Nama Group of Namibia,
These fossils are still under study in ilar to Praecambridium from Ediacara. mentioned above as implying a young
connection with problems of annelid- These assemblages, irrespective of sedi- Ediacarian age.
arthropod relations. Two primitive ar- mentary facies, share the numerical The poor showing in the Phanerozoic
thropods, Praecambridium and Parvan- dominance of the phylum Cnidaria, re- fossil record of soft-bodied Metazoa,
corina, are rare. Together with the enig- flecting evolutionary level at Ediacarian abundant in the present marine fauna, is
matic triradiate Tribrachidium (resem- time. From some regions (central Aus- probably due to preservational or col-
bling edrioasteroid echinoderms without tralia, China, Scandinavia, and North lecting bias, the evolution of macropha-
calcareous plates) they amount to but a Carolina), however, finds reported are gous predators, an increase in sapropha-
few percent of the fauna (Table 1). limited or unique. gous macro- and microbiota, or some
788 SCIENCE, VOL. 217

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combination of such factors. Biostrati- be done in paleoichnology, but what has chaeocyathids or a bioherm builder) are
graphic studies have shown that the been observed confirms the rapid mor- unknown below the Nemakit-Daldyn
"sudden" appearance of abundant small phological and ethological diversification and lower Baltic beds of the Soviet
shells of the Tommotian Stage (basal of the Metazoa during the transition from Union, widely considered to be upper
Cambrian) was preceded in the latest Ediacarian to Cambrian, a product of Yudomian (Ediacarian) in Siberia but
Ediacarian (latest Yudomian of Siberia their increasing exploitation of the troph-post-Vend (post-Ediacarian) in Europe-
and probably the Sinian of the Yangtze ic resources of the sea floor and its an Russia. Elsewhere we have not seen it
Platform) by the occurrence of three or sediments in the littoral and neritic below the basal Cambrian (Tommotian),
four genera of minute tubular and spicu- zones. which leads us to regard the Epiphyton-
lar fossils (Anabarites trisulcatus, proba- bearing beds of the Nemakit-Daldyn as
bly a worm tube; Protohertzina, similar Cambrian. A fifth algal type, the large,
to chaetognath grasping spicules; and Plant Microfossils and Stromatolites usually compressed, algal spheroids that
Cambrotubulus and/or Hyolithellus). comprise the form genus Chuaria,
The seemingly abrupt appearance of Paleobotanical evidence related to the ranges through the uppermost Protero-
small shelly fossils in many places is age, paleoecology, and correlation of zoic, from strata perhaps as old as 1200
enhanced by the common transgression Ediacarian rocks comes from microscop- million years or more to its very top.
of Early Cambrian on older sediments, ic filamentous algae, acritarchs (small, Here it overlaps the lower range of Bav-
leaving gaps in the record. Ecological spheroidal, organic bodies of unknown linella and may extend into the Paleozoic
effects of this transgression are well dis-but probably plant origin), stromato- under other names.
played in England (55). The onset of lites (accretionary buildups, mainly of Vidal (57) is more explicit with refer-
biomineralization was part of the rapid CaCO3 and thought to be generally of ence to Eurasian acritarchs. Among 22
early Phanerozoic diversification of the blue-green algal origin), and perhaps previously known acritarch taxa he stud-
Metazoa. No universally causal extrinsic some ambiguous structures called micro- ied, he finds that 18 percent first appear
environmental factor has been credibly phytolites. in the Vend, 10 percent are limited to the
invoked for it. The microbial flora of the pre-Phaner-Vend, and 6 percent are limited to the
Metazoan diversification during the ozoic and early Phanerozoic sediments is upper Riphean (uppermost Proterozoic).
Ediacarian-Cambrian transition was par- only beginning to receive the attention it We are not sufficiently confident about
alleled by an increase in diversity of warrants. Available evidence, however, either the identification of acritarch spe-
trace fossils (3, 53, 56). These are indica-implies that such a flora has existed from cies or the ages of the stratigraphic units
tors of mainly metazoan life activities on Archean times to the present and that discussed (for instance, the upper Sinian
and in sediments, made by live animals forms preserved increased markedly in Suberathem of northern China) to accept
at the places where their markings are size, abundance, and diversity during these estimates as a basis for the defini-
found. They may reveal much about the later Proterozoic and early Phanerozoic tion of systemic boundaries. We are
nature of these activities (feeding, loco- time. Although such material has not yet hopeful, however, that continued acri-
motion, and so on), but only exceptional- been systematically studied in the Edia- tarch research will eventually lead to a
ly do they indicate the place of the carian and immediately underlying strata better Proterozoic biostratigraphy and
originator in the zoological classifica- of Australia, it is known to be biostrati- contribute to a more cogent definition
tion. Different kinds of traces can be graphically useful in correlative beds than we can presently give for the base
made by different activities of the same elsewhere and will become more so as of the Ediacarian.
animal and similar traces by different our now limited knowledge of it in- Stromatolites undergo a decline in
animals. Thus the number of named form creases (57). abundance and diversity with the transi-
genera does not indicate taxonomic di- The ranges of four distinctive, strati- tion from Proterozoic to Phanerozoic,
versity but ethological differentiation. graphically limited, organic-walled, mi- and with the appearance of metazoan
Known Ediacarian trace fossils, prob- crobial forms known to us are shown on browsers, eukaryotic precipitation and
ably of detritus feeders, are simple shal- the left of the correlation chart. Spiny secretion of CaCO3, and competitive ex-
low burrows and relatively poorly orient-planktonic acritarchs of the sort called clusion. They are, however, still rela-
ed search trails on bedding surfaces. ByMicrhystridium (58) and the spiral fila- tively abundant during Ediacarian time.
contrast, early Cambrian assemblages ments of the microalga Obruchevella Although few are uniquely Ediacarian,
include deep, vertical burrows (Sko- (59), illustrated in Fig. 4, apparently first
an association of certain distinctive co-
lithos), complex burrows (Phycodes pe- appear in strata of Ediacarian age and lumnar forms with strongly enveloping
dum, Diplocraterion, Chondrites, Trep- range upward through the Cambrian. laminae or lateral wall-like structures
tichnus, and Plagiogmus), excavations The spheroidal, multicomponent form would suggest a late Proterozoic to Edia-
made by trilobite-like arthropodan ap- genus Bavlinella, representing the endo- carian age (61). The simple peripherally
pendages (Rusophycus), and a variety of sporangia or clonal colonies of Sphaero- enveloped columns of Boxonia grumu-
trackways. Fedonkin (53) recognized congregus (57, 60), ranges from upper losa, combined with branching columnar
Ediacarian trace fossils as dominantly Proterozoic through Ediacarian equiva- "walled" forms like Gymnosolen and
two-dimensional and horizontal, in con- lents into the Cambrian of the Northern the lumpy Linella ukka and Paniscol-
trast to three-dimensional Early Cambri- Hemisphere. Vidal (57) describes it as a lenia, would be such an association.
an forms. Some relatively complex loco- characteristically Vendian form, but re- Where found above a sequence contain-
motion trails that occur only rarely in the cords it in the Cambrian and in and ing evidence of extensive late Proterozo-
Ediacarian become common in the Cam- beneath the Varangerian tillites, as well ic glaciation an Ediacarian age would
brian, for instance, Didymaulichnus, as in other truly upper Proterozoic rocks seem likely.
which suggests the actions of a mollusk- such as those of the upper Sinian "Su- Finally, in the Soviet Union and to
like gliding foot rather than the peristal- berathem" of northern China. Bushlike some extent in China, subdivision and
tic movement of a wormlike animal. colonies of the bifid branching microalga correlation of late Proterozoic and early
Much more analytical work remains to Epiphyton (commonly commensal on ar- Phanerozoic strata has, in some in-
27 AUGUST 1982 789

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 Plants ITracefossils Soft-body fossils Shelly fauna N. Central
Eg -a0 E South Central Southwest Southern Siberia Russian Scandinavia Midland Sou
E gE~ a co=Australia
'O Australia
o Africa
c (Amadeus China (Composite platformpafr
? (Namib
(Composite (Composite England New
c5 - - a - ECEpoch
=~~~~~~ (Flinders .~ =(E.Yangtze Anabar- (Composite) N. Sweden- (Nuneaton
= - - =. Anabar-
0 l E o -= O a (Sri Ranges) Basin) Desert) Gorges) O eS. Norway) Leices
ve=0 aomo 0
- S
a E 0
- Eo -a- e
5 li , li.0E
a0 .- 0 . u
i0 Cii ~Sip zU0- ll (52,79, and
- =_ o E . 0- ? 0. (30,31) (77) (29) (71,78) other (53,70,80) (40,81) (37,38,
maS
5 W~C Ol 0~~i-.'3O.2C
0 ui Q n00:3 0. o: in co > O
F- o0 00 (C
< 0 -? a. < B O..Ia<,,sources)
< ^sucs
4^~ 4~ 4~ 4~ A44~._- Todd River Tian- Tyussver
Todd RiverVergale Strenuaeva
. .ss._. Brigus-Brigus
/~'FmF
Dol. Fish heban Fm. Fm. 4 Shale Stocking
Hawker River Fm. Lukati Smithpper fo
- Early Group , TC Upper ford P
Early GpP p Sub- Shipal - (Talsy) Sandstone Sha
~ (Lower)r
C(Lower) Para-group Fm.
Arumbera cuPestro-
Iuangsha-Fm.
tsvet
Camb- china . dong Fm. Fm. Zone.T arnhl
E- ~ ~ ~ .n 5Lontova. 7 Hartshil
' ? "* ~ ~~~~~~ Uratanna Tc c LMelmatu
/0 a/// Neai-
U. MNmsd eimatu TC FmTc
o ~~~ ~~Fm. T0 IT 14 Daidyn. c ~_ - Vangsg
N a . >.tanMem.
ssO?-*~ A hiban-d Fm--/-
Rovno p
Fm F
o . "'cwaz c :C SChwa
_ Turkut
~~~~~~~~~~~~~~~~~~o Caldecote
0Goupas
Hamadingl (Star-ay
\^ 'Xexmatu
' Late ///////////////Lae Rand o Mem R/h - Kotln . , K ^ VolcanI
? - - (Upper) ^ ^ ^^^LSubgrpt Mem.
Por,. P~ Rehka
ma- (Kanilov) ullingitf,$ /Con
a c Edia- /// /// //o m Brand
- carian // Kuibis* Doushantuo K Grou
I- i awns^ Lower ?aSub- PR Fm' CKhatspit
U Sub~Bonney sC *C s P T
______ Y 0 --rumbera
Arumbera* -0 O ./?' M
G

~~~~i~~~~~~i a~~~~~~~s
? I ^ <? group 4)IMaple-
^"^ *C/^ ^</^7///
ill ~~~~~~~~~~~ ~~~Early groupFm Maastakh > w
X. S CPDT o0 ^^
~... . hia(Lower) FWonoka Fm. Fuii F m.
0 Edia- BunyerooFm. peRedkinora G Fm. ?

w carianA g tataka E
Brachina Fm. C ////// E
* Edia~~~~~~~~~~~~~carlia n rocksy////~/'~..
Fm. 0?~.. Ekre
)W~ ? 7 ~ ~ Nuccaleena F. Cap Dol / Shale 0
670 A AVA /ICF Av4
4 Trilobites A Evidence
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-n'- - of glaciation Olymp. G _ r e
^~~~~~~~~~~~~~~~~~~.
Tc Cambrian r u'~ u~m:u670?htsi approximate Later /A A////// //// Vicha tillite n ?o
traceo~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'
fossils 550?, g///
T Simple tracks age in years x 106 Proterozoic N//////uo/ Gariep Nnu Group B
t>J ~ ~ ~ ~ ~ ~ ~ ~ ~ ' ru~, n
FCribricyathida
^J ~ ~ ~ ~ ~ ~ ~ ~m.
~ ~ ~ Formation (commonly Umbertana Aralka roup Fm. Lower Fm.o
~~ Cribricyathida ~ ~ ~ ~ ~ws oe~~or
u F. with evidence Gop F. (Numees 'Lobroo
J Small Cambrian shelly fossils Oz. Quartzite Gu Fm.) Liantuo Fm. zat - Rihe Hedm
c3 Ediacarian medusoids Ss. Sandstone of ice action) Areyonga R ivpanvv Sse Riphean Group
Ediacarian soft-body fossils, Dol. Dolomite V A Fm.A Crystalline System
subscript denotes taxon Mem. Member G rocks

Fig. 5. Provisional global correlation of Ediacarian System. (N

z
5

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stances, been based on a heterogeneous
group of microstructures referred to as
microphytolites. Although we have
doubts about the biological nature of
most of these structures and the validity
of subdivision and correlation based on
them, several named entities have been
reported from Ediacarian and adjacent
strata in South Australia, and it is
claimed that partial success in correla-
tion was attained in tests based on sam-
ples of provenance unknown to the iden-
tifier (62).
Global Distribution
Beyond South Australia, Ediacarian
fossils are known from the Amadeus
Basin of central Australia and from some
dozen other regions on four other conti- B. S. Sokolov, 23d Int. Geol. Congr. Contrib.
13.
nents (Africa, Asia, Europe, and North
America). Figure 5 summarizes the main
features of nine of these fossiliferous
sequences plus two western North
American sequences that are expected
eventually to yield Ediacarian fossils.
Space, regrettably, does not permit fur-
ther discussion of these areas in this
article. Their stratigraphy, biotas, and
paleoecology, however, are detailed in
references cited at the top of each re-
gional column.
It should be stressed that Fig. 5 shows
only relative positions in geologic se-
quence and does not conform to any
scale of thickness or time, other than as
ages are indicated for the top and base of
the Ediacarian. The underlying upper
Proterozoic is simply whatever occurs
beneath the Ediacarian, with no special
reference to age other than the supposed
approximate time equivalence of the
probably glacial rocks. The overlying
Lower Cambrian here includes only the
Tommotian and Atdabanian Stages of
earliest Cambrian age.
In addition to sequences shown in Fig. northern China reference section for the Sinian
5, evidence available is consistent with
an Ediacarian presence in North Caroli-
na in the United States (47); the upper
Brioverian of Brittany, Normandy, and
the Channel Islands (63); Antarctica (64);
South America (65); and the northern
Arabian Shield (59). Also possible is an
Ediacarian presence somewhere beneath
or in the basal part of the Candana
Quartzite of northern Spain (66). In In-
dia, Svalbard, and Alaska also, well-
preserved sedimentary rocks of the rightpart of the Moscow basin, where it was studied
age range and facies to yield an Ediacar-
ian biota may be present. Indeed, there
is reason to believe that future discovery
will show the Ediacarian to have an areal
extent worthy of its rich historical and
evolutionary interest (85).
27 AUGUST 1982
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References and Notes and other reasons, Keller (74), Yakobson and
Krylov (75), and other Soviet geologists have
1. R. C. Sprigg, Trans. R. Soc. South Aust. 71, 212
(1947).
deplored the acceptance of the "greater Ven-
dian." If, as many believe, the late Proterozoic
2. M. F. Glaessner, ibid. 81, 185 (1958); Nature
glaciogenic deposits of the East European Plat-
(London) 183, 1472 (1959); Sci. Am. 204
(March), 72 (1961); Biol. Rev. Cambridge Phi-
form and the margins of the Baltic Shield are of
approximately the same age as those of Austra-
los. Soc. 37, 467 (1962).
lia, only the Valdai (including Redkino and
3. , Lethaia 2, 369 (1969).
Kotlin strata) would correspond to the Ediacar-
4. , Geol. Rundsch. 47, 522 (1958).
ian as here defined, not the official Vendian
5. __ and B. Daily, South Aust. Mus. Rec. 13,
sensu lato which Sokolov accepts at least since
369 (1959); M. F. Glaessner and M. Wade, Pa-
1973 (52). Vendian has also been used in this
laeontology 9, 599 (1966).
broad sense by Harland and Herod (76) and by
6. L. C. Noakes, 20th Int. Geol. Congr. El Sistema
Cambrico (1956), part 2, pp. 213-238.
Cowie and Cribb (34), among others, to include
both fossiliferous rocks of Ediacarian age and
7. R. Richter, Senckenbergiana Lethaea 36, 243
the Varangerian tillites beneath.
(1955).
8. T. D. Ford, Proc. Yorks. Geol. Soc. 31, 211
19. M. Wade, Trans. R. Soc. South Aust. 94, 87
(1970).
(1958).
20. R. J. F. Jenkins, Geol. Soc. Aust. Abstr. 1st
9. G. H. Chadwick, Geol. Soc. Am. Bull. 41, 47
Geol. Conv. (1975), p. 21.
(1930).
21. M. R. Walter, personal communication to P.C.,
10. H. E. Wheeler, J. Geol. 55, 153 (1947); C. J.
who examined the fossils and agrees to their
Stubblefield, 20th Int. Geol. Congr. El Sistema
metazoan origin.
Cambrico (1956), part 1, pp. 1-44; V. V. Mis-
22. W. B. Harland and I. N. Herod, Geol. J. Spec.
sarzhevsky and A. Yu. Rozanov, Izv. Akad.
Issue 6, 189 (1975).
Nauk SSSR Ser. Geol. 2, 60 (1963); P. Cloud,
23. P. R. Dunn et al., Nature (London) 231, 498
Evolution 2, 322 (1948).
(1971).
11. M. F. Glaessner, J. Geol. Soc. Aust. 10, 223
(1963).
24. R. J. F. Jenkins, Geol. Soc. Aust. Abstr. 1st
Aust. Geol. Conv. (1975), p. 19; R. J. F. Jen-
12. H. Termier and G. Termier, Rev. Gen. Sci.
kins, P. S. Plummer, K. C. Moriarty, Trans. R.
Pures Appl. Bull. Assoc. Fr. Av. Sci. 67, 79
(1960).
Soc. South Aust. 105, 67 (1981).
25. The enigmatic structure called Bunyerichnus is
so far known only from the fragment shown in
Sov. Geol. Probl. 9 (1968), pp. 5-15; Geol.
Geofiz. 12 (No. 6), 13 (1971); 24th Int. Geol. Fig. 2F, its counterpart, and one smaller frag-
Congr. Sec. 1 (1972), pp. 78-84; 23d Int. Geol. ment. The view illustrated is of an upper bed-
Congr. Rec. Int. Paleontol. Union (1972), pp. ding surface. The arcuate grooves and ridges of
this specimen, from the scalloped outer margin
79-84; Paleontol. Zh. 1, 3 (1976).
to the faint groove at the lower edge (x in Fig.
14. B. S. Sokolov, Geol. Geofiz. 15 (No. 2), 3
(1974).
2F) deviate only slightly from a set of concentric
circles that would close on themselves, if contin-
15. V. E. Savitsky, Geol. Mag. 115, 127 (1978); A.
ued, with a maximum diameter of 24 cm. They
Yu. Rozanov, ibid. 104, 415 (1967); A. Yu.
Rozanov et al., The Tommotian Stage and the clearly represent either (i) the tracing of a long-
ish (12 cm) stalklike, flexible structure of some
Problem of the Cambrian Lower Boundary,
complexity that swung by currents about a point
(Doklady Geologiya Instituta Akademii Nauk
of attachment or (ii) the imprint of a large (24 cm
555R 206, Moscow, 1969); A. Martinsson, Leth-
in diameter) discoidal structure, having enough
aia 13, 46 (1980); M. F. Glaessner, Bull. Geol.
Soc. Am. 82, 509 (1971); J. W. Cowie and M. F.
plasticity to account for asymmetries observed.
These characteristics imply that it was the prod-
Glaessner, Earth Sci. Rev. 11, 209 (1975); J. W.
uct of a once living organism, very likely a
Cowie and A. Yu. Rozanov, Geol. Mag. 111,
237 (1974).
metazoan. Although such a marking might be
produced by the swinging about a central point
16. P. Cloud, in Evolution and Environment, E. T.
Drake, Ed. (Yale Univ. Press, New Haven,
of an elongate wormlike form [for instance,
Conn., 1968), p. 37; Paleobiology 2, 351 (1976).Vermiforma Cloud (in 47)] or a pennatulacean
(53), the positive epi-relief displayed is more
17. B. S. Sokolov, Geol. Geofiz. 15 (No. 4), 18
consistent with it being an imprint than a drag
(1974); Paleontol. Zh. 1, 3 (1976).
18. The terms "Eocambrian" (67, 68) and "Infra- mark; for example, compare Fig. 2F with Fig.
2B. The counterpart imprint also shows features
cambrian" (68, 69) are ambiguous. They are not
resembling gonadal imprints of some recent
based on biological criteria, no type sequences
medusae. We conclude that Bunyerichnus is
have been designated, and they have been used
certainly biologic, very probably metazoan, and
loosely to include widely varying parts of the
Precambrian sequence and locally, though unin-
most likely medusoid.
tentionally, rocks now known to be Cambrian 26. P. Cloud, L. B. Gustafson, J. A. L. Watson,
Science 210, 1013 (1980).
(for instance, the Ringsaker Quartzite of Nor-
way, upper member of the Vangsas Formation).
27. J. W. Durham, Annu. Rev. Earth Planet. Sci. 6,
They have been generally abandoned. Sinian, 21 (1978); V. M. Palij, Dopov. Akad. Nauk Ukr.
RSR Ser. B 36 (No. 1), 34 (1974).
although once officially adopted in the Soviet
Union for rocks of Vendian and Riphean age
27a.H. Faul, personal communication to P.C.
28. H. D. Hedberg, Ed., International Stratigraph-
(70), suffers similar disadvantages. It is still
widely used in China, but in two very different
ic Guide (Wiley, New York, 1976), pp. 66-93.
29. G. J. B. Germs, Bull. Precamb. Res. Unit Univ.
senses. The thin Sinian System of the Yangtze
Gorges and neighboring parts of southern China, Capetown 12, 1 (1972); Am. J. Sci. 272, 752
younger than about 680 to 720 million years (71)
(1972); A. Kroner, M. D. Williams, G. J. B.
Germs, A. B. Reid, K. E. L. Schalk, ibid. 280,
and with glacial sediments at its base, is appar-
942 (1980).
ently not even represented in the standard
29a.R. Lacour-Gayet, A Concise History of Austra-
Suberathem. Only if the glacial sediments were lia, J. Grieve, Transl. (Penguin, Blackburn, Vic-
toria, Australia, 1976), p. 3.
excluded from its base and the local basal Cam-
brian (Tommotian) from its top would it approxi- 30. W. V. Preiss, R. W. R. Rutland, B. Murrell, in
mately coincide in time with our Ediacarian Precambrian of the Southern Hemisphere,
D. R. Hunter, Ed. (Elsevier, Amsterdam, 1981),
System.
The term most deserving consideration and pp. 327-354.
comparison with the Ediacarian is Vend or 31. R. Goldring and C. N. Curnow, J. Geol. Soc.
Vendian. This name was proposed by Sokolov Aust. 14, 195 (1967); B. G. Forbes, Trans. R.
(72), who later (73) referred to the Ediacarian as Soc. South Aust. 95, 219 (1971); P. S. Plummer,
"the Australian equivalent of the Vendian." ibid. 102, 25 (1978).
This equivalence is less straightforward than it32. Briefly, the divisions of the Wilpena Group,
may appear. A new stratigraphic scale for the comprising the type Ediacarian System in Bun-
Proterozoic of the Soviet Union was accepted yeroo Gorge, are (from bottom upward) as fol-
by a large number of specialists at a conference lows:
in Ufa in 1977 (74). This makes the Vend the 1) Nuccaleena Formation; basal Wilpena
uppermost division of the Proterozoic, including Group, conformable on the underlying locally
(from base upward) the Vilchan, Volyn, and tillitic rocks of the Umberatana Group. Pinkish,
Valdai Groups. Its stratotype is in the western laminated, flaggy dolomite going upward to pur-
ple shale with dolomite lenses. These were the
from numerous bore holes. It is inaccessible to initial shallow marine sediments of the postgla-
direct observation. cial warming and marine innundation that intro-
The Vend was originally equated with the duced the Ediacarian System. Thickness, 10 m.
Valdai only (72). It was later extended down- 2) Brachina Formation; reddish-brown to ol-
ward to include the Volyn, partly volcanogenic ive-green, thin-bedded, micaceous siltites with
(70), and then the Vilchan, partly glaciogenic shale and fine-grained sandstone. Bunyerichnus
(together comprising the Drevlyany "series" of near the middle. Current marks, flute marks,
some). The Ediacarian fauna of soft-bodied Me- local coarsening and grading, and ripple marks
tazoa is found only in the Valdai, and, for this imply submarine conditions of varying depth,
791
All use subject to http://about.jstor.org/terms
 with flat-topped ripples indicating local subaeri- 38. P. J. Patchett, N. H. Gale, R. Goodwin, M. J. shida, Nature (London) 122, 466 (1969); E. L.
al exposure. Thickness, - 1200 m. Humm, J. Geol. Soc. London 137, 649 (1980). Yochelson and H. E. Herrera, Ameghiniana 11,
3) ABC Range Quartzite; light colored, flaggy 39. R. P. Coates and W. V. Preiss, Precamb. Res. 283 (1974); T. R. Fairchild, Soc. Bras. Geol.
to massive, cross-bedded and ripple-marked, 13, 181 (1980). 30th Congr. Bras. Geol. Recife Bol. 1, 181
ridge-forming, locally feldspathic sandstone and40. 0. Kulling, Sver. Geol. Unders. Ser. Ba 19, 28 (1978).
quartzite. Flat-topped ripple marks and desicca- (1964). 66. T. P. Crimes, J. Legg, A. Marcos, M. Arboleya,
tion polygons indicate episodes of subaerial ex- 41. J. R. Pringle, Geol. Mag. 109, 465 (1973). in Trace Fossils 2, T. P. Crimes and J. C.
posure within a shoaling marine environment. 42. N. M. Chumakov, Dokl. Akad. Nauk SSSR 198 Harper, Eds. (Seel House, Liverpool, 1977), pp.
Thickness in Bunyeroo Gorge is 80 to 120 m, (No. 2), 419 (1971). 91-138.
thickening westward. 43. C. J. Hughes and W. D. Bruckner, Can. J. 67. W. C. Br0gger, Norge i 19de Aarhundred (Kris-
4) Bunyeroo Formation; monotonous, red- Earth Sci. 8, 899 (1971). tiana Central Torylekerret, 1900), vol. 1, pp. 1-
dish-brown and green, silty shales with thin 44. H. W. Fairbain, S. Moorbath, A. 0. Ramo, 32; B. Asklund, Colloq. Int. CNRS 76, 39
cupriferous dolomite bands, local carbonaceous W. H. Pinson, P. M. Hurley, Earth Planet. Sci. (1958).
shale, and dolomitic sandstones showing local Lett. 2, 321 (1967). 68. R. B. Neuman and A. R. Palmer, 20th Int. Geol.
ball-and-pillow structure and sole markings in- 45. S. B. Misra, Bull. Geol. Soc. Am. 80, 2133 Congr. El Sistema Cambrico (1956), part 2, p.
dicative of a western source and gravity mass (1969). 427.
transport. Basally equivalent Elkera Formation 46. S. Bengston and T. P. Fletcher, U.S. Geol. 69. N. Menchikoff, Bull. Soc. Geol. Fr. 16, 451
of central Australia contains metazoan trace Surv. Open-File Rep. 81-743 (1981), p. 18; B. (1949); P. Pruvost, Bull. Geol. Soc. Belge 60, 43
fossils (21). Thickness in Bunyeroo Gorge is Greene and H. Williams, Can. J. Earth Sci. 11, (1951).
- 400 m (reaching 700 m elsewhere). 319 (1973). 70. B. S. Sokolov, Colloq. Int. CNRS 76, 104
5) Wonoka Formation; greenish-gray calcare- 47. P. Cloud, J. Wright, L. Glover III, Am. Sci. 64, (1958).
ous siltstones and fine-grained channel sand- 396 (1976). 71. F.-T. Chung, Sci. Sin. 20 (No. 6), 818 (1977).
stones in the lower part, going upward to mainly 48. R. J. F. Jenkins and J. G. Gehling, Rec. South 72. B. S. Sokolov, Izv. Akad. Nauk SSSR, Ser.
cross-bedded, silty, commonly sole-marked, de- Aust. Mus. 17 (No. 23), 347 (1978). Geol. 5, 21 (1952).
trital limestones, locally stromatolitic. Thick- 49. M. F. Glaessner, in Treatise on Invertebrate 73. , Siberian Branch Akad. Nauk SSSR
ness is 460 m in Bunyeroo Gorge, but the Paleontology (Geological Society of America, Trans. Inst. Geol. Geofiz. 232, 5 (1975).
formation thickens dramatically, changes facies, Boulder, Colo., 1979), part A, p. 79; and 74. B. M. Keller, Geol. Mag. 116, 419 (1979).
and cuts downward across underlying se- M. R. Walter, in Precambrian of the Southern 75. K. E. Yakobson and N. S. Krylov, Int. Geol.
quences in a series of submarine canyon depos- Hemisphere, D. R. Hunter, Ed. (Elsevier, Am- Rev. 20, 709 (1978).
its dispersed south to north across the center of sterdam, 1981), pp. 361-396. 76. W. B. Harland and I. N. Herod, Geol. J. 6, 189
the Copley map sheet, next north of the Para- 50. H. D. Pflug, Palaeontogr. Abt. A. 134 (Nos. 4- (1975), special issue.
chilna sheet [C. C. von der Borch, R. Smit, 6), 226 (1970); G. J. B. Germs, Lethaia 6, 1 77. M. R. Walter, Aust. Bur. Miner. Res. Geol.
A. E. Grady, Flinders Univ. Inst. Aust. Geo- (1973). Geophys. Rep. 214 (1980), microform Mf92 with
dyn. 81/3 (1981), pp. 1-44]. 51. L. I. Borovikov, Dokl. Akad. Nauk SSSR 231 chart.
6) Pound Subgroup; massive ridge- and bluff- (No. 5), 1182 (1976). 78. Y. Qian, M.-E. Chen, Y.-Y. Chen, Acta Paleon-
forming sandstone and quartzite, including the
52. B. S. Sokolov, Mem. Am. Assoc. Pet. Geol. 19, tol. Sin. 18 (No. 3), 207 (1979); R-J. Cao and Y-
lower Bonney Sandstone and upper Rawnsley 204 (1973). Z. Liang, Acad. Sin. Mem. Nanjing Inst. Geol.
Quartzite: 53. M. A. Fedonkin, Tr. Geol. Inst. Akad. Nauk Paleontol. 5, 1 (1974); P. Cloud, field notes,
6A) Bonney Sandstone; mostly arkosic and SSSR 342, 1 (1981). October 1979; M. F. Glaessner, field notes, May
micaceous lenticular siltstones showing ripple 54. C. R. Scotese et al., J. Geol. 87, 217 (1979); 1979.
marks, desiccation polygons, local ball-and-pil- P. A. Jell, ibid. 82,319 (1974); L. P. Zonenshain 79. S. N. Serebryakov, personal communication.
low structure, and cross-bedding, including and A. M. Gorodnitskiy, Geotektonika 2, 3 80. A. P. Brangulis, V. V. Kirjyjanov, K. A. Mens,
trough cross-beds indicative of fluviatile pro- (1977). A. Yu. Rozanov, U.S. Geol. Surv. Open-File
cesses. A periodically emerged coastal environ- 55. M. D. Brasier, R. A. Hewitt, C. J. Brasier, Rep. 81-743 (1981), p. 26; B. S. Sokolov, per-
ment is indicated. Thickness, - 300 m. Geol. Mag. 115, 21 (1978); M. D. Brasier and sonal communications.
6B) Rawnsley Quartzite; whitish quartzite R. A. Hewitt, Palaeogeogr. Palaeoclimatol. 81. S. Foyn and M. F. Glaessner, Nor. Geol.
and sandstone with local siltite layers, locally Palaeoecol. 27, 35 (1979). Tidsskr. 59, 25 (1979); K. Bj0rlykke, A. Elvs-
ripple-marked and cross-bedded, gradational 56. S. P. Alpert, in Trace Fossils 2, T. P. Crimes borg, T. Hoy, ibid. 56, 233 (1976); K. Bj0rlykke,
from Bonney Sandstone beneath. Contains t '- and J. C. Harper, Eds. (Seel House, Liverpool, Geol. Surv. Can. Pap. 78-13 (1978), p. 49.
cal Ediacarian body fossils and imprints a 1977), pp. 1-8; N. L. Banks, in Trace Fossils, T. 82. M. M. Anderson, Can. J. Earth Sci. 9, 1710
massive, basal, bluff-forming unit. Represents P. Crimes and J. C. Harper, Eds. (Seel House, (1972); B. Greene and H. Williams, ibid. 11, 319
final overfilling of Adelaide Geosyncline prior to Liverpool, 1970), pp. 19-34; J. W. Cowie and (1974); S. Bengston and T. P. Fletcher, U.S.
late Ediacarian and Cambrian truncation and A. M. Spencer, in ibid., pp. 91-100; B. Daily, Geol. Surv. Open-File Rep. 81-743 (1981), p. 18.
onlap of initial Cambrian seas. Thickness, Univ. Adelaide Cent. Precamb. Res. Spec. Pap. 83. F. G. Young, Geol. Surv. Can. Bull. 288, 1
- 500 m. Unconformably overlain by argilla- 1 (1972), p. 13; B. D. Webby, Lethaia 3, 79 (1979); H. Gabrielse, Am. J. Sci. 272, 521 (1972).
ceous sandstones of Parachilna Formation, (1970). 84. C. A. Nelson, Geol. Mag. 115, 121 (1978); P.
Hawker Group (lower Cambrian), with vertical 57. G. Vidal, Fossils Strata 9, 1-57 (1976); Nor. Cloud, Am. J. Sci. 273, 193 (1973); S. P. Alpert,
trace fossils Skolithos and Diplocraterion. Geol. Unders. Bull. 362, 1 (1981). in Trace Fossils 2, T. P. Crimes and J. C.
Total thickness, - 2970 m. 58. W. A. S. Sarjeant, Rev. Micropaleontol. 9, 201 Harper, Eds. (Seel House, Liverpool, 1977),
The description above applies to the strato- (1967); S.-C. C. Lo, Precamb. Res. 13, 109 vol. 2, pp. 1-8.
type section and nearby outcrops. The sequence (1980). 85. While this article was in press a parallel endorse-
is broadly similar over other parts of the Flin- 59. P. Cloud, S. M. Awramik, K. Morrison, D. G. ment of a new basal Phanerozoic System and
ders Ranges but with the variations in thickness Hadley, Precamb. Res. 10, 73 (1979). Period was published by R. J. F. Jenkins [Trans.
and facies to be expected in an evolving mobile 60. M. Moorman, J. Paleontol. 48, 524 (1974). R. Soc. South Aust. 105 (No. 4), 179 (1981)].
shelf environment (30). 61. P. E. Cloud and M. A. Semikhatov, Am. J. Sci. 86. We are grateful to so many individuals and
33. K. A. Plumb, J. H. Shergold, M. Z. Stefanski, 267, 1017 (1969); M. R. Walter, Stromatolites organizations for help with aspects of the re-
B.M.R. J. Aust. Geol. Geophys. 1, 51 (1976). and the Biostratigraphy of the Australian Pre- search reported here that space does not permit
34. J. W. Cowie and S. J. Cribb, in The Geologic cambrian and Cambrian (special paper 11, Pale- appropriate acknowledgment. We mention
Time Scale, G. V. Cohee, M. F. Glaessner, ontological Association, London, 1972); M. R. therefore only our special obligation to Dr. W.
H. D. Hedberg, Eds. (American Association ofWalter and W. V. Preiss, 24th Int. Geol. Congr. V. Preiss of the South Australian Department of
Petroleum Geologists, Tulsa, Okla., 1978), pp. Sec. 1, 85 (1972). Mines and Energy, to the several colleagues in
355-362. 62. W. V. Preiss and I. N. Krylov, Izv. Akad. Nauk the Soviet Union and China who furthered our
35. L. Rose, personal communication to M.F.G. SSSR Geol. Ser. 7, 61 (1980). researches there, and to the Queen's Fellow-
36. S. J. Cribb, J. Geol. Soc. London 131, 203 63. A. D. Squire, Geol. Mag. 10, 223 (1973). ship Committee of the Australian Department of
(1975). 64. E. L. Yochelson and E. Stump, J. Paleontol. Science and the Baas Becking Geobiological
37. M. D. Brasier and R. A. Hewitt, U.S. Geol. 51, 872 (1977). Laboratory, who sponsored P.C.'s year in Aus-
Surv. Open-File Rep. 81-743 (1981), p. 29. 65. C. A. L. Isotta, A. C. Rocha-Campos, R. Yo- tralia.

792 SCIENCE, VOL. 217

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