Journal ofthe Geological Society, London, Vol. 149, 1992, pp. 607-613, 6 figs.

Printed in Northern Ireland

Vendobionta and Psammocorallia: lost constructions of Precambrian evolution

A. SEILACHER
Geologisches Institut der Universitat Tiibingen, Sigwartstr. 10, 7400 Tiibingen, Germany and Kline Geology Laboratory,
Yale University, POB 6666, New Haven, CT 06511, USA

Abstract: The non-availability of biomineralized skeletons and lowlevels of predation led Vendian evolu-
tion along strange avenues. The Ediacara-type Vendobionta appear to represent a in kingdom,
which foliate
shapes, large sizes and the necessary compartmentalization were achievedby quilting of the skin rather than
by multicellularity.Psammocorallia,incontrast,areinterpreted as coelenterates that constructed an
internal sand skeleton. Both were immobile soft-bottom dwellers that had high population densities, and
both became preserved by obrutional accidents; thus they render ‘fossil snap shots’, in which the original
distributional patterns, age structures and standing biomasses of populations are accurately recorded.

With the exception ofstromatolites,a small cone-shaped
calcareous fossil of problematic affiliation (Cloudinu) and a
number of small organic-walled fossils, known Precambrian
organisms were largely soft-bodied. How the general lack of
biomineralization was related to the chemistry of the Precam-
brian ocean is a matter of discussion, but means that the bio-
logical sink for carbonatewas in any case rather different from
that in Phanerozoic oceans. On the other hand, preservational
‘snapshots’ of communities suggest that the standing biomass
was not significantly lower than in equivalent modern environ-
ments, at least with regard to the epibenthic realm.
On the basis of constructional and preservational features,
threekinds of Vendian macro-fossils can be distinguished:
Vendobionta, trace fossils and sand corals (Psammocorallia).
Vendobionta
As pointed out earlier (Seilacher 1988), the interpretation of
Ediacara-type organisms as soft-bodied ancestors of modern
metazoan phyla (Glaessner 1984) meets with difficulties.
Rather, they are considered to be a unique, quilted type of
biological construction that has no counterpartin the modern,
or even the Phanerozoic, biosphere. A joint TiibingenNale
expedition to Newfoundland in the summer of 1990 (Mark
Brandon, chief investigator) hasyielded additional evidence in
support of this unorthodox view. In order to emphasize the
basic difference, the informal name Vendozoa is here changed
to Vendobionta with the following definition.
Kingdom Vendobionta. Immobile foliate organisms of diverse geo-
metries that were only a few millimetres thick, but reached several
decimetres in size. A shared characteristic is the serial or fractal quilt-
ing of the flexible body wall, which stabilized shape, maximized exter-
nal surface and compartmentalized the living content.
Since no organs can be recognized, this content is thought to have
been a Plasmodial fluid rather than multicellular tissue. Included are
the Petalonamae (Pflug 1972) and a variety of forms previously inter-
preted as soft-bodied ancestors of metazoan phyla. R a n g e : Vendian.
ClaimedCambriansurvivors seem to showdifferentpreservational
properties.
Nature of the quilted construction
While overall geometries and symmetries vary agreatdeal
(Fig. l), quilting is the primarysharedcharacteristic of all
607
Vendobionta. As the organisms grew, either new quilts were
inserted marginally or existing unitsexpanded and became
subdivided fractally. Both modes have the result that quilts do
not exceed acertaindiameter.Thisconstruction limits the
available morphospace in three ways. (1) Quilts never bundle,
but are added in one plane to make foliate bodies that may
reach almost one metre in size but are only a few millimetres
thick. (2) The quilt pattern and relief was the same on either
side of the carpet, as can be verified in specimens that became
partly folded-over before being cast (Fig. 3). (3) Individual
quilts never protrude beyond the common margin to form
branches, tentacles or appendages.
Function of quilting
The quilted hydrostatic constructionallowed the Vendobionta
to maintain a foliate morphology that deviated considerably
from a simple balloon shape. Such bodies could, like carpets,
rest firmly on the sea floor. At the same time they facilitated
direct metabolic exchange with the environment by their
maximized surface-to-volume ratio. If Vendobionta were not
multicellular, the size limits set by Runnegar (1982) would not
apply, since diffusion could be assisted by flow processes.
At the same time quilting may have served an internal func-
tion. Plants and animals developed multicellularity to over-
come the size limits set for unicellular organisms. In other
words: they use the originally reproductive process of cell
division for growth purposes. Subdividing a syncytial proto-
plasm by quilting appears to be a valid alternative, which has
its analogues in the skeletal chamberlets and canal systems of
giant foraminifera, or the quiltedumbrella of Acetubuluriu. So,
primitive organisms could growto large sizes by integumental
compartmentalization without necessarily becoming multicel-
lular; but this development had consequences. As Gould
(1989) haspointed out, quiltedcreaturescould never have
reached the degree of tissue differentiation required for higher
levels of organization. In addition, they were much more vul-
nerable than multicellular organisms. A predator could not
only swallow the whole organism, but even small bites would
probably have led to fatal leakage from the living bags. Thus
the disappearance of the Vendobionta at, orbefore, the evolu-
tionary radiation of metazoans with hard jaws and claws is not
surprising. Even if the seapen-like fossils fromthe Burgess
Shale (Conway Morris 19896, fig. sa) were indeed vendobion-
tan survivors, the picture hardly changes: no such impressions
608 A . SEILACHER

LIFE STYLES OF BILATERAL

VENDOBIONTA dh

Ernietta Pferidinium Dickinronia Phyllozoon spriggins

Fig. 1. As immobile organisms, the Vendobionta depended on sedimentation rates. In areas of low sedimentation they could recline as flat
carpets or stand erect, provided that the basal disc or ring sufficed as an anchor in the soft sediment. In areas of increased background
sedimentation, the margins grew up to form bag-like sediment stickers. Note that serial and fractal modes of quilting are represented in all
three modes of life. Forms from Mistaken Point, Newfoundland (lower right) are not yet formally described.

have ever been found in lower Cambrian sandstones that are
taphonomically and environmentally more equivalent to the
Ediacara deposits.
Life styles
Whether they lived photo- or chemosymbiotically or simply
absorbed dissolved organicsubstances,there is little doubt
that Vendobionta were immobile. Even though they were pri-
marysoft-bottom dwellers, they shared adaptationalprob-
lems, and strategies to solve them, with the secondary soft-
bottom dwellers derived from sessile animal stocks of later
times (corals, oysters, brachiopods, cirripeds). The vast major-
ity of Vendobionta, including leaf-shaped forms, lived as
carpet-like recliners in areas where rates of background sedi-
mentation were low enough not to cover them. In Namibia,
however, the margins of variously quilted carpets grew up-
wards with sand sedimentation and thustransformed into bag-
like sediment stickers, the way they have been preserved in the
sediment (Fig. 1). They may also have become passively re-
implanted after scouring,if the ‘rock-in-the-sock’ model of the
Psammocorallia (Fig. 4) is modified into one filled with loose
sand.
A special problem are pennate forms with an ‘attachment
disk’ at their base. In soft sediment, such a disk would be a
much poorer anchorthan root-like extensions. One could
argue that vendobiontan construction did not allow for the
fabrication of such extensions. Alternatively, one might con-
sider these structures to be floats rather than anchors. The latter
idea would appear particularly suitable in the case of unde-
scribed Churniu-like forms from Newfoundland (Fig. 3),
whose bases resemble a life-belt or a doughnut rather than
being concentrically quilted. Still, biostratinomic evidence
falsifies this model. While associated species maintain their
original random orientations, the stalked species are always
current-aligned with the disks pointing upcurrent. So the rings
did function as anchors, although the exact mechanism is not
yet clear. From their mode of preservation, it appears that the
rings were fluid-filled, like the quilted leaf part. On the other
hand, the ring tube far exceeds the diameter allowed for in-
dividual quiltings in the foliate parts, suggesting different
physiological constraints. Is it possible that such a structure
could also effect scour-implantation, even though its content
was not much denser than the water around?
Another problem is the stalk of the Newfoundland ‘sea-
pens’. Not only isit much morepronouncedthan in true
 VENDOBIONTAANDPSAMMOCORALLIA 609

Fig. 2. Impressions of various Vendobionta buried under an ash layer (chiselled edge in lower part of picture) at Mistaken Point,
Newfoundland. Note that only stalked forms are current-orientated, probably by felling rather than transport. Later, the impressions became
tectonically deformed, roughly in the direction-of the pre-existing slope. Diameter of coin atleft hand margin is 1.7cm.

Fig. 3. Ashfall produced a downslope

current, the direction of which is
recorded by the felling direction of
stalked species, the scour in front of a
folded-over leaf-shaped form and
cross-bedding in the ash layer. Another
current direction is recorded by bent
tops of stalked species and folding-over
of spindle-shaped forms, which also
show that top and bottom surfaces
looked alike. The block diagram
explains relief inversion in the stem
parts, in which spindle-shaped
organisms that came to lie under the
stem are also involved. All drawings
made from oriented silicon casts made
at Mistaken Point, Newfoundland.
610 A . SEILACHER

Charnia, but it also lacks the sharp contours of rings and leaf preservational events.(4) The mere abundanceof Vendobionta
parts and is always preserved with oppositerelief. Presumably, on Vendian sea floors, in addition to low levels of predation
the stalk was filled with a more solid, gel-like material-the and scavenging, madetheir episodic preservationlikely inspite
best such an organism could do to produce a rigid support of an extremely low fossilization potential.
structure. This material would have taken more time to disinte-
grateand consequentlywouldhaveproducedtheinverted
collapse relief, without sharp outlines (Figs 2-3), that charac-
Vendian trace fossils
terizes the stalk portion in all Newfoundland specimens. In contrast to the Vendobionta, the makers of Vendian trace
Becoming erect rather than reclining would be alright in a fossils were true, worm-like animals. In order to plough or
photosymbiotic organism, but does not agree with bacterial burrow through the sediment, an organism needs the co-ordi-
chemosymbiosis at the sediment-water interface. On the other nated action of appendages or cilia, or the ability to move by
hand, perfectlypreservedVendobiontahave been found in undulation or peristalsis, as is typical for metazoans. On the
NorthCarolina in avolcano-turbidite facies together with other hand, trace fossils share with the Vendobionta an auto-
flysch-typeichnofossils
(Gibson 1989), indicating
depths chthonouspreservation,althoughthey do notprovidetrue
below the photic zone. So the trophic modeof the Vendobionta snapshots, because traces recorded on one bedding plane are
remains an open question. Possibly they were more ambivalent not necessarily contemporaneous.
in this respect than multicellular organisms. Stratigraphical changesof trace fossil faunas across the Pre-
cambrian-Cambrian boundary in many sections of the world
have been described by Crimes (1987, 1989, this volume) and
Preservational conditions Fedonkin (19906). These workers found a general upward in-
Vendobionta are always preserved as mere impressions withoutcrease in diversity, and listed a considerable number of ich-
any organic substance left behind. Otherwise they occur in very nogenera that are either restricted to the Vendian or continue
differentsedimentologicalsituations: atthe bases of storm into thePhanerozoic,as well asa still largernumber that
sands in South Australia, below graded ash layers in New- appear in the early Cambrian. Still, a lot needs to be done in
foundland, with turbidites in North Carolina,
orwith determining synonymies among these taxa and in relating them
supposedly estuarine inunditesin Namibia. What thesesettings to particular modesof burrowing and feeding strategies, and to
have in common is the role of episodic event sedimentation, by coelomate, acoelomate and pseudocoelomate body construc-
which the biota became accidentally buried. It has been argued tions.
that the absence or low level of bioturbation on Precambrian While increasing diversity meets the expectation, the etho-
sea bottoms favoured the preservation of these delicate impres- logical character of Precambrian trace fossils does not. If the
sions. I would argue that the mere abundance of Vendobionta radial burrow systems that have been variously described as
lying around on the sea floorwas an equally important factor. medusae (Cloud & Glaessner 1982; Fedonkin 1990a, pl. 6, fig.
In the Mistaken Point Formation of Newfoundland, the 5) are added to the guided meanders, there is a surprising com-
fossils (about 9 species of Vendobionta plus large ‘medusoid’ plexity ofbehaviour patterns, reminiscentof those that charac-
blobs of problematic origin) occur only on bedding planes thatterize deep sea ichnocoenosesin the Phanerozoic. Such trends
had originally been covered by graded layers of volcanic ash may in fact have begun muchearlier. Large radial impressions
(Fig. 2). Cross-bedding, discovered by F. Pfluger in the coarser in the early Proterozoic Notenius Formation of northern Aus-
bottom parts of these layers, suggests that the ashfalls pro- tralia (Robertson 1960) have been discarded as sand volcanoes
duced downslope currents, with a southward direction of flow (Walter 1972). Asconfirmed by S. Dzulynski (pers. comm.
that agrees well with the alignment of the stalked ‘sea pens’. 1991), however, the published pictures look much too regular
Oncethetectonicstrainhasbeeneliminated,otherVend- for an inorganic origin and resemble radial feeding burrows. So
obionta (including the common spindle-shaped formand leaf- the question remains, whether low levelsof competition or
shaped forms without a basal disk) show random orientations. similartrophicconditions (bacterial mats)madesediment
Except for incipient scouring along the upcurrent of edges
some feeders in Precambrian shallow seas behave similarlyto those
Vendobionta, there areno erosion marks. Nordid we observe on deep-sea bottoms at later times.
any current-swept accumulations of fossils. Thus it is safe to But are all occurrences of Vendobionta shallow marine?
assume that the current was weak and did not transport the Greaterdepthhas been suggested for theMistakenPoint
carpets before burial. Formation of Newfoundland(ConwayMorris 1989~).The
Sudden obrution appears to apply also to the fossil hori- chief argument is the absence of wave-induced features (oscil-
zons in Namibia. Blocks in thecollection of H. D. Pflug lation ripples; hummocky cross stratification) and grading in
(Giessen) show closely packed bag-like organisms in 3-dimen- the ash layers, which could occurat any depth. Since there are
sional preservation (Pflug1973). They are never collapsed, and no trace fossils in this part of the section, the exact palaeo-
always maintaintheir original upright life positions, indicating bathymetry remains ambiguous.
that they were partly sand-filled during life. Similar bag-like Considering this background, the localities in the Slate Belt
forms have also been found in USSR (Fedonkin 1990a, pl. 5 , of NorthCarolinagainimportance. So far, onlya few
figs. 1,4, 5,6;PI. 8, fig. 5 ; PI. 10, figs 2-3; pl. 12, figs 3, 5 ; pl. 16, specimens of Pteridinium have been found which undoubtedly
figs I , 3, 6). belong to the Vendobionta; but associated are delicate trace
Four conclusions can be drawn ( l ) Vendobionta were im- fossils (Gibson 1989). Among these is a very distinctive form
mobile epibenthic organisms. (2) They are likely to occur as that has been attributed to Syringomorpha, but which is better
‘fossil snapshots’, in which population and age density, popu- assigned to Oldhamia. The same species hasbeen found in the
lation structure and standing biomass at the dayof the eventare PuncaviscanaFormation of Argentina(Aceiiolaza 1979),
registered without the usual taphonomic distortion. (3) These where it is associated with Oldhamia radiata and Nereites, and
values can be extrapolated to intervening time slices, in which also with trilobite tracks, in a more typical flysch facies.
these organisms failed to leave a record due to the absence of The North Carolinaconnection thus re-opens the argument
 VENDOBIONTAANDPSAMMOCORALLIA
as to whether Oldhamia is also Vendian in other occurrences
(Alaska, New York, Argentina, Ireland, Belgium), or whether
it spans the Precambrian-Cambrian boundary. Secondly, the
palaeobathymetry of these deposits becomes relevant for the
biology (photosymbiotic or not) of the Vendobionta.
Psammocorallia
Placing the Vendian medusoids of earlier workers in either the
Vendobionta or trace fossils, we are faced with another diffi-
culty. While bilaterian ancestors are recorded by trace fossils,
where are the roots of the equally basic group of coelenterates?
In addition, DNA sequencing (D. Bridge, Yale, pers. comm.)
suggests that Anthozoa, rather thanScyphozoa, were the most
primitive coelenterates.
The claim now is that corals are in fact represented in the
Vendian biota, although in an unfamiliar guise (Seilacher
1990). Some of the putative medusoids are distinguished from
others because of a different kind of preservation. They are
perfectly circular in outline and either protrude fromsole faces
as hemispherical bodies with an unusually high relief (e.g.
Nemiana; Fedonkin 1990a, pl. 5, fig. 3) or they occur on top
surfaces as sharply delineated positive epireliefs with irregu-
larly spaced concentric rings (e.g. Tirasiana).
The interpretationof these fossils is not derived from Vend-
ian material, but frombetter known representatives in the
Mickwitzia Sandstone (lower Cambrian, Sweden; Figs &6)
andtheConulariaSandstone (upperOrdovician, Jordan;
Seilacher 1983). Here, they are found in tempestite beds and
always in the same top-and-bottomorientation. Asmooth
hemispherical base, sometimes with a central dimple, resem-
bles assumed actinian burrows (Eergaueria). In the Swedish
material, the upper surfaceof the bodies has a reticulate sculp-
ture made of somewhat finer sand (Protolyella;Fig. 4), or it is
finely pitted and bears a variable number(3-5) of strong radial
ridges (Spatangopis; Fig. 5). The Jordanian specimens have
narrower and more numerous radial ridges as well as heavy
concentric rings that continue in cross section into hemi-
spherical growth laminae (Seilacher 1983).
61 1
Fig. 4. The internal and organically
cemented sand skeleton of
Psammocorallia served mainly as an
anchor that currents and waves
automatically implanted into the sandy
substrate. In Protolyella, growth laminae
were added mainly on the smooth
bottom surface covered by ectoderm. On
the flat top (specimens in the collection
of Jan Johansson), the digesting
endodermal surface was increased, not
by radial septa, but by a reticulate
sculpture. In tempestitic slabs
(Riksmuseet, Stockholm), animals were
washed together in life-orientations
along the troughs between spill-over
ripples, where the muddy part of storm
sedimentation has cast not only the
skeletons, but also marks of the rocking
implantation process. Mickwitzia
Sandstone, Sweden. Scale bar is 1 cm.
The traditional view, that these are scyphozoan medusae
whose gastral cavity became sand-filled after death, is unten-
able. Not only would it imply that the medusae were all
embedded in upside-down positions, but theradial mesenteries
should appear as recesses, rather than ridges, in a cast. Mor-
phology and internal structureare inconsistent with them
being trace fossils (Jensen 1991), though the somewhat similar
Erooksella fromthe middle Cambrian of Alabama (Harr-
ington & Moore 1956) is actually a radial burrow system.
In the new interpretation, these fossils are regarded as inter-
nal sand skeletons of coelenterates comparable to actinians.
They were built of sand grains that entered the gastral cavity
and then became phagocytized and deposited in place of a
mesogloea between ecto- and endoderm. Originally, this endo-
skeleton was rigid, since closely-packed fossils never indent
each other. Nevertheless, soonafter burial worms could
burrow right throughthem and compactioncoulddeform
them. Thus the cement of these sand skeletons was probably
organic and after death became readily digested by microbial
activity.
Functionally, such construction resembles that of solitary
soft-bottom corals. The massive skeleton provided the im-
mobile animal with an anchor, notonly by weight but also by
its shape. As conceived in the ‘rock-in-the-sock’ model (Fig. 4),
such an organism would become automatically implanted into
a sandy bottom by current or wave scour. In addition, the
skeleton could be used to increase the area of the digesting
endodermalsurface by developing reticulate, concentric or
radial rugosities.
On the other hand, there are inherent differences between
these and the external calcareous skeletons of later corals. As
an internal structure, the sand skeleton did not grow into a
cone-shaped theca, into which the soft parts could withdraw,
nor could it be cemented to hard substrates, Thus there is no
attachment scar even in the smallest individuals. Also the
septa are broad and blunt, in contrast to calcareous septa,
whose sharp edges are dictated by sphaerolithic biomineraliz-
ation in small blisters indenting the large body pneu.
The strange construction of sand corals also had important
612 SEILACHER A.
preservational consequences. In their sandy habitats, fossiliz-
ation potential was normally minimal because the skeleton
disintegrated into loose sand before it could leave a lasting
impression. Only if instantaneously embedded in fine mud
could the shapes of these skeletons be preserved. This is why
they are foundonly in sandy tempestites, either with the round
bottoms sticking out from thesole face, or as complete bodies
on rippled top surfaces, where they became blanketed by the
muddy tail of ,storm sedimentation.Thus, as in the Vend-
obionta, event-induced obrution provides us with fossil
snapshots, in which original populations are portrayed
directly.
The ubiquity of Psammocorallia on sandy bottoms of the
lower Cambrian of the Baltic Province is also suggested by
another phenomenon. Early stratigraphers coined the name
Eophyton Sandstone. Today, everybody agrees that Eophyton
has nothing to do with algae, but represents tool marks made
by objectsdragging over themud. Tool marks are regular
features on the sole faces of sandy tempestites deposited in later
times; but they usually have the shapes of impact casts made by
bouncing shells and shell fragments. The continuous bands of
Eophyton are unique and can be used as time markers in spite of
being pseudofossils. Therefore it is here suggested that sand
skeletons, skipped over the ground by storm waves, were in-
volved in the production of Eophyton.
In theMickwitzia Sandstone, sandskeletons occur in a vari-
ety of shapes that are asyet only partly appreciated. In addition
to Spatungopsis and Protolyellu, there are ear- and fan-like
forms, some of which bear delicate, bryozoan-like reticulation
patterns (Fig. 6). Others look like colonies of Fuvosites.
Associated small cones of Volborthellu represent still another
class of sand skeletons. Possibly there was aplethora of
unrelated animal groups, including sponges, that all used sand
instead of biomineralization to make rigid skeletons. But they
also fell under the same preservational constraints.
Thus it is surprising that nothing similar has as yet been
described from elsewhere in the lower Cambrian. The many
specimens of Brooksellu fromthe middle Cambrian of
Tennessee resemble Spatungopsis and have likewise been inter-
Fig. 5. In Spatangopsis, the bottom
surface still has the smooth and
rounded bag shape, sometimes with a
central dimple. On the endodermal top,
however, the surface is not only pitted,
but also bears 4-5 heavy and blunt
radial septa that variously deform the
round outline. In the extreme case (top
right) the whole skeleton becomes
propeller-shaped, with the ectodermal
surface being reduced to a small pad
(specimens from Jan Johansson
collection). In the slab (Riksmuseet,
Stockholm), impressions of tentacles are
also preserved. Mickwitzia Sandstone,
Sweden. Scale bars are 1 cm.
preted as gastric casts of medusae (Harrington & Moore 1956).
However, they are preserved as ironstone concretions and are
actually radial spreite burrowscomparableto Teichichnus
stellutus. Thus, identification of the Palaeozoic Psammocoral-
lia with similar Vendian fossils still needs to be verified.
In any case, the Psammocorallia add another radiation to
the history of anthozoans, predating that of the calcitic Tabu-
lata and Rugosa and the much laterone of the aragonitic
Scleractinia. The following definition is proposed.
Psarnrnocorallia. Radially symmetrical sand bodies, with a smooth,
hemisperical lower surface, commonly with a central dimple. Upper
surface flat or with a variable number of prominent radial septa and/or
concentric growth rings, as well as more delicate ornamentation (reti-
culate or tuberculate). In vertical section, surficial growth rings may be
seen to continue intohemispherical sand layers parallel to the bottom
surface.
These fossilshave traditionally beendescribed ascastsof scy-
phozoan stomachs. Theyare here interpreted as internal sand skeletons
that stabilized anthozoan polyps living on sandy bottoms. Since these
skeletons lost their organic cement soon after death, they could be
preserved only when smothered by mudfall after a storm, almost in-
variably in life position. Range: Vendian to Ordovician. Typicalgenus:
Protolyella.
Other genera: Spatangopsis and various Vendian
‘Medusoids’.
Conclusions
(1) Megascopic Vendian organisms are difficult to place in
modern phyla, classes and orders. Rather, they appear to rep-
resent independent radiations that ultimately could not com-
pete in metazoan evolution.
(2) All known groups were benthic, with a shallow, in-
fauna1 tier of worm-like bilaterian sediment feeders (trace
fossil record) and immobile primary soft-bottom dwellers at
the sediment surface (Vendobionta; Psammocorallia). Thus,
the Cambrian revolution implied probably not only the radia-
tion of biomineralizing metazoan phyla, but also a general
mobilization and an expansion of the zone inhabited by mega-
scopic organisms deeper into the sediment and into the open
 VENDOBIONTAANDPSAMMOCORALLIA 613

Fig. 6. Other fossils with the same

preservational charactistics, but of
different shapes, are associated with
typical Psammocorallia in the
Mickwitzia Sandstone (lower Cambrian,
Sweden). It is as yet impossible to
decide whether they are also sand corals
or members of other phyla (e.g.
sponges) that used a similar mode of
making skeletons. Two fan-shaped
bodies in the bottom centre are from the
Riksmuseet, Stockholm; the other
specimens are in the Johansson
collection in Skollersta (Sweden). Scale
bars are 1 cm.

water. To test this assumption, one should determine whether FEWNKIN, M. A. 1 9 9 0 ~ .SystematicdescriptionofVendianMetazoa. In:
or not Vendian black shales actually contain remains of nek- SOKOLOV,B. S. & IWANOWSKI, A.
B.
(eds) The Vendian Sysrem, 1.
Springer Verlag, 71-120.
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(3) Within the epibenthic zone, standing biomass was sur- IWANOWSKI, A. B. (eds) The Vendian Sysrem, 1. Springer Verlag,
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Received I I April 1991; revised typescript accepted 15 December 1991