Discovery of the oldest bilaterian from the Ediacaran
of South Australia
Scott D. Evansa,1,2, Ian V. Hughesb, James G. Gehlingc, and Mary L. Drosera
a
Department of Earth Sciences, University of California, Riverside, CA 92521; bSection of Ecology, Behavior and Evolution, Division of Biological Sciences,
University of California San Diego, La Jolla, CA 92093; and cDepartment of Palaeontology, South Australia Museum, Adelaide, SA 5000, Australia
Edited by Neil H. Shubin, University of Chicago, Chicago, IL, and approved February 17, 2020 (received for review January 21, 2020)
Analysis of modern animals and Ediacaran trace fossils predicts
that the oldest bilaterians were simple and small. Such organisms
would be difficult to recognize in the fossil record, but should have
been part of the Ediacara Biota, the earliest preserved macro-
scopic, complex animal communities. Here, we describe Ikaria
wariootia gen. et sp. nov. from the Ediacara Member, South Australia,
a small, simple organism with anterior/posterior differentiation.
We find that the size and morphology of Ikaria match predictions
for the progenitor of the trace fossil Helminthoidichnites—indica-
tive of mobility and sediment displacement. In the Ediacara Member,
Helminthoidichnites occurs stratigraphically below classic Ediacara
body fossils. Together, these suggest that Ikaria represents one of
the oldest total group bilaterians identified from South Australia,
with little deviation from the characters predicted for their last com-
mon ancestor. Further, these trace fossils persist into the Phanerozoic,
providing a critical link between Ediacaran and Cambrian animals.
bilaterian | Ediacaran | Ediacara Biota | phylogenetics | trace fossil
T he first macroscopic animal fossils are recognized within the
soft-bodied Ediacara Biota (1, 2). Among these are candi-
date poriferans (3), cnidarians (4), and ctenophores (5). Rare
Ediacaran taxa have been interpreted as putative bilaterians,
namely, Kimberella (6, 7). However, small furrowed trace fossils
are generally accepted as definitive evidence for total group
bilaterians in the Ediacaran (8–10). The size and morphology of
these trace fossils suggest that they were produced by millimeter-
scale organisms that would be difficult to recognize in the fossil
record (11).
Helminthoidichnites are horizontal trace fossils found in Edia-
caran and Phanerozoic deposits globally (12, 13). Helminthoi-
dichnites is a curvilinear burrow that can be preserved on both
bed tops as well as bottoms and occurs most commonly on the
base of thin (submillimeter to millimeter scale) discontinuous
sand bodies, or shims (8, 14). The preservation of Helminthoi-
dichnites in negative relief flanked by positive levees on bed
bottoms indicates that the progenitor moved under thin sand
bodies following deposition and burial, displacing sediment (8,
9, 11, 14). Observed relationships between intersecting Hel-
minthoidichnites indicates the ability of the progenitor to move
vertically, albeit on millimeter scales (11). Rare Helminthoidichnites
penetrating body fossils of macroscopic taxa may represent the
oldest evidence of scavenging (11).
In modern environments, Helminthoidichnites-type structures
can be produced by a variety of bilaterians (9, 11). A likely pro-
genitor for Ediacaran Helminthoidichnites has yet to be identified,
although it has been suggested that these were produced by simple
“worm-like animals” (9). Critically, based on the nature of sedi-
ment displacement by a horizontally burrowing organism, it would
have been small, with a maximum diameter less than that observed
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for Helminthoidichnites. Such behavior necessitates anterior–pos-
terior differentiation, as well as a coelom, consistent with bilaterian-
grade tissue organization (8, 9, 11, 15).
Helminthoidichnites are preserved abundantly within the
Ediacara Member, Rawnsley Quartzite in the Flinders Ranges
and surrounding regions of South Australia (16). The Ediacara
www.pnas.org/cgi/doi/10.1073/pnas.2001045117
Member consists of shallow marine sandstone event beds 50 to
500 m below a basal Cambrian disconformity (17). At the National
Heritage Nilpena site, the excavation and reconstruction of 37-m-
scale fossiliferous bed surfaces reveals in situ communities of the
Ediacara Biota (18). At Nilpena, and sections within the Flinders
Ranges, Helminthoidichnites occurs more than 100 m below the
first appearance of Kimberella (19, 20). There are currently no
radiometric dates to constrain the absolute age of the Ediacara
Member; however, significant overlap of taxa with well-established
deposits from the White Sea region of Russia indicates that these
are likely between 560 and 551 million years old (21–24). A similar
pattern of leveed, horizontal trace fossils (although in this case
DEVELOPMENTAL
assigned to the ichnogenus Archaeonassa) occurring strati-
graphically below classic White Sea assemblage body fossils in
BIOLOGY
Russia (9, 23) may corroborate the early appearance of trace
fossils in South Australia.
Results
AND PLANETARY SCIENCES
Here, we report the discovery of the new genus, new species
EARTH, ATMOSPHERIC,
Ikaria wariootia, the interpreted progenitor of Helminthoidichn-
ites. We have identified 108 Ikaria on a single bed surface (1T-A)
and 19 from float at multiple localities, preserved in negative
hyporelief on the base of sandstone beds (Fig. 1). Ikaria is found
in fine-grained sandstones in two facies representing deposition
in relatively shallow marine environments between fair-weather
and storm-wave base (14, 17, 25).
Significance
The transition from simple, microscopic forms to the abundance
of complex animal life that exists today is recorded within soft-
bodied fossils of the Ediacara Biota (571 to 539 Ma). Perhaps
most critically is the first appearance of bilaterians—animals
with two openings and a through-gut—during this interval.
Current understanding of the fossil record limits definitive evi-
dence for Ediacaran bilaterians to trace fossils and enigmatic
body fossils. Here, we describe the fossil Ikaria wariootia, one of
the oldest bilaterians identified from South Australia. This or-
ganism is consistent with predictions based on modern animal
phylogenetics that the last ancestor of all bilaterians was simple
and small and represents a rare link between the Ediacaran and
the subsequent record of animal life.
Author contributions: S.D.E., J.G.G., and M.L.D. designed research; S.D.E., I.V.H., and M.L.D.
performed research; S.D.E., I.V.H., J.G.G., and M.L.D. analyzed data; and S.D.E., I.V.H.,
J.G.G., and M.L.D. wrote the paper.
The authors declare no competing interest.
This article is a PNAS Direct Submission.
Published under the PNAS license.
1
Present address: Department of Paleobiology, Smithsonian Institution, Washington,
DC 20560.
2
To whom correspondence may be addressed. Email: EvansSD@si.edu.
This article contains supporting information online at https://www.pnas.org/lookup/suppl/
doi:10.1073/pnas.2001045117/-/DCSupplemental.
PNAS Latest Articles | 1 of 6
 A B
Fig. 1. Type specimen of I. wariootia from Nilpena, including (A) photograph; and (B–D) 3D laser scans. Notice distinct bilateral symmetry (wider end
identified by white star in C and deeper end by black star in D). P57685. (Scale bars, 1 mm.)
Systematic Description
Ikaria wariootia gen. et sp. nov.
Etymology. The generic name is after the word “Ikara,” which is the
Adnyamathanha name for Wilpena Pound, and means “meeting
place” in the Adnyamathanha language. Ikara is the major land-
mark in view from Nilpena, and the fossil has been named to
acknowledge the original custodians of the land; species are
named for Warioota Creek, which runs from the Flinders Ranges
to Nilpena Station.
Holotype. P57685 (Fig. 1; South Australia Museum).
Paratype. P57686 (Fig. 2A; South Australia Museum).
Field Paratypes. 1T-A bed 001 to 007 (Fig. 2 B–J; Nilpena).
Horizon and Locality. Ediacara Member, Rawnsley Quartzite at
the National Heritage Nilpena field site and Bathtub Creek.
Diagnosis. Irregular millimeter-scale ovoid preserved in negative
hyporelief. The major axis length averages 2.3 times the minor
axis. There is distinct asymmetry along the major axis with one
end wider and more broadly curved (white star in Figs. 1C and 2
D–F, G, and J). In profile, the broader end is preserved in more
significant negative relief and with a steeper curvature (black star
in Figs. 1D and 2 H and I). Rare specimens are bent about the
long axis (Fig. 2 F and J) and/or exhibit potential evidence of
modularity, with two to five body divisions (Fig. 2 D and E).
Description. I. wariootia are well-defined elongate ovals, fusiform
in shape (Figs. 1 and 2). Three-dimensional (3D) laser scans
demonstrate clear anterior/posterior differentiation, with one
end distinctly smaller and more tightly curved. Length of the
major axis ranges from 1.9 to 6.7 mm and the minor axis from 1.1
to 2.4 mm. Preserved depth ranges from 0.6 to 1.6 mm. There is a
consistent linear relationship between total length and total
width (SI Appendix, Fig. S1A). The relationships between total
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length and depth (SI Appendix, Fig. S1B) as well as width and
depth (SI Appendix, Fig. S2) are irregular. Depth is always less
than width, suggesting that fossils of Ikaria are compressed. This
confirms previous interpretations that the preserved depth of
specimens from the Ediacara Member is strongly influenced by
taphonomic processes (e.g., ref. 26).
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C D
While the morphology of Ikaria is very simple, it is consistent
across specimens and is unambiguously distinct from other
structures. The consistent shape and length-to-width ratio are
not what is observed for rip-up clasts of organic mats, which are
irregular (14). Although mat rip-ups are found within the
Ediacara Member, they do not occur in the same lithologies and
facies as Ikaria (14, 25), which represent deposition in a lower-
energy environment. Furthermore, rip-up clasts have a different
biostratinomic and diagenetic history than Ikaria and all other
body fossils (14). The outer margin of Ikaria is sharp, and they
are preserved with considerable relief, distinct from the sur-
rounding matrix and organic mat textures (Fig. 2). This is con-
sistent with other nonsessile taxa from the Ediacara Member
(27), suggesting that Ikaria represents the body fossil of a free-
living organism.
Ikaria can be easily differentiated from other taxa preserved on
the same bed surface and of similar size and scale (SI Appendix,
Fig. S3). Thus, it is unlikely a juvenile form of a previously de-
scribed taxon. The lack of larger specimens with comparable
morphology suggests that maximum size is ∼7 mm. The recogni-
tion of other taxa on the same surface preserved at the same scale
and with similarly well-defined outer margins distinct from the
organic mat corroborates the biologic, body-fossil origin of Ikaria.
Specimens of Ikaria are found in association with Helmin-
thoidichnites, albeit rarely (Fig. 2A). The range of Ikaria widths
plots entirely within those measured for Helminthoidichnites with
the maximum size of body fossils not exceeding that of trace
fossils (SI Appendix, Fig. S4). Further, the Anderson–Darling test
indicates that size-frequency distributions are not significantly
different (P value 0.448). This, combined with clear anterior–
posterior differentiation, suggests that Ikaria is the only known
contemporaneous body fossil with the suite of characters pre-
dicted for the progenitor of Helminthoidichnites.
Discussion
In general, it is rare to have trace fossils and the organisms that
produced them preserved together, particularly with respect to mo-
bile metazoans. This can be attributed to both the different preser-
vational pathways between body and trace fossils and the ability of
the animal to move away from the area where it left evidence of
activity (28, 29). In certain cases, the morphological characteristics of
body fossils from the same deposits can be used to reliably determine
the progenitors of particular trace fossils (e.g., ref. 30).
Body fossils in the Ediacara Member, including Ikaria, are well
preserved on the bottoms of centimeter-scale sandstone beds
Evans et al.
 A

B C

DEVELOPMENTAL
BIOLOGY
D E F

AND PLANETARY SCIENCES

EARTH, ATMOSPHERIC,
G I J

H
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Fig. 2. Photographs (A and B) and 3D laser scans (C–J) of I. wariootia. (A) Specimen (white arrow) associated with Helminthoidchnites. (B–E) Associated
specimens; black boxes in B and C are the same specimen shown in close up in negative hyporelief (D) and inverted (E). (F and J) Bent specimens. (G and H) N
bedding plane (G) and profile (H) of the same specimen. (I) Profile demonstrating variable relief. Notice correlation between broader, wider end (white stars)
in the bedding-pane view and more significant relief end (black stars) in the profile. (A) P57686. (B–E) 1T-A 001 to 003. (F ) 1T-A 004. (G and H) 1T-A 005. (I)
1T-A 006. (J) 1T-A 007. (Scale bars, 1 mm.)

Evans et al. PNAS Latest Articles | 3 of 6

 with early mineralization of overlying sand casting the tops of
these organisms following burial (8, 14, 16). Although counter-
parts are identified in rare cases on bed tops, these are poorly
preserved and at a resolution that is unlikely to produce identi-
fiable features at the same scale as Ikaria. In contrast, Helmin-
thoidichnites is found on both bed tops and bottoms, but most
commonly on the base of millimeter-thick shims, where well
preserved body fossils are rare (8, 14). Negative hyporelief
preservation indicates that Helminthoidichnites formed after the
deposition of overlying sand, with the organism that produced it
capable of moving into and out of thin layers of sand (11). This
predicts that we should only find Helminthoidichnites and its
progenitor on the same bed bottom in rare instances when it died
while burrowing underneath thin sand bodies. Given the simple
morphology and preservation of both body and trace fossil in
negative relief, even if Ikaria was preserved at the end of a trail, it
is unlikely that it would be possible to confidently identify as
distinct from that trace. We interpret the surprising discovery of
Helminthoidichnites with nearby Ikaria (Fig. 2A) as the result of
vertical movement from the bedding plane in the region between
the end of its trace fossil and its final resting place. While this
scenario was likely exceedingly rare, it may represent the only
situation in which it would be possible to distinguish associated
body and trace fossils and further corroborates interpretations of
Ikaria as the progenitor of Helminthoidichnites.
We propose that Ikaria is the trace maker of Helminthoi-
dichnites and potentially the oldest, definitive bilaterian, at least
as represented in the fossil record of South Australia. Kimberella,
the only other taxon from the Ediacara Member that is consis-
tently reconstructed as a bilaterian, occurs significantly higher
stratigraphically than the earliest appearance of Helminthoi-
dichnites (6, 7, 19, 20). Similarities between taxonomic assem-
blages have been consistently cited as evidence that White Sea
assemblage fossils from the Ediacara Member, including Kim-
berella, are conservatively 560 to 551 Ma (21–24). The strati-
graphic position of Helminthoidichnites suggests that the first
appearance of Ikaria was likely within this age range or possibly
earlier. Burrows initially interpreted to be from much older
Ediacaran rocks in Uruguay have uncertain age constraints (31,
32). Trace fossils from Brazil, representing the activity of
meiofaunal bilaterians, occur 30 to 40 m above a tuff dated at
555 Ma and in close association with Cloudina, indicating that
they are likely younger than Ikaria (10, 24). A recently described
segmented bilaterian from South China, associated with trace
fossils, is interpreted to be younger, larger, and more complex
than Ikaria (33).
The ability to move and produce recognizable trace fossils is
not unique to bilaterians. Complex body and trace fossils from
older Ediacaran deposits were probably produced by muscular
eumetazoans interpreted to be cnidarians (34). Dickinsonia, and
similar Ediacara Biota fossils, likely do not represent crown-
group bilaterians, but were mobile and left trace fossils (35,
36). Modern protists generate simple burrows, but are typically
smaller than Ikaria (37, 38). Laboratory experiments demon-
strate that mobile foraminifera form burrows in clay and silt;
however, they do not produce burrows in fine-grained or coarser
sand (37). Large testate amoeba in deep-sea environments are
associated with horizontal trails similar to those observed in the
Ediacaran, but these are surficial and represent movement by
rolling (38). Flatworms are mobile, but do not burrow below the
sediment–water interface and rarely leave trace fossils (39).
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Among these examples, expression on bed bottoms with furrows
is unique to Helminthoidichnites and suggests mobility associated
with significant displacement of medium sand grains. This is
consistent with reconstructions of Ikaria containing musculature
and a coelom (15, 40). Combined with the relative size of body
and trace fossils, these characteristics are unique to bilaterians.
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Fig. 3. Reconstruction of Ikaria in life position forming a Helminthoidichnites-
type trail.
Polarity of relief and curvature characterize anterior/posterior
differentiation in Ikaria (Fig. 3), supported by directed move-
ment in trace fossils. Preservation of v-shaped transverse ridges
within Helminthoidichnites suggests peristaltic mobility (ref. 11;
Fig. 2A). Ikaria morphology implies a potentially modular body
construction, which would have aided in muscular organization
required for peristalsis (40). Sediment displacement and scav-
enging reveal that Ikaria likely had a coelom, mouth, anus, and
through-gut (11, 15, 40), although these are unlikely to be
reproduced in the fossil record. Preferential preservation of
Helminthoidichnites under thin sand bodies indicates that Ikaria
sought out these environments, possibly due to increased oxygen
availability (11, 14, 20). Ultimately, as the depth of overlying sand
increases, oxygenated environments give way to sulfidic, in-
hospitable settings due to decomposition of organic matter, sup-
ported by the restriction of Helminthoidichnites to beds <15 mm
thick (11, 14, 20). Ikaria was likely able to detect organic matter
buried in well-oxygenated environments as well as potentially toxic
conditions, suggesting rudimentary sensory abilities. Combined,
these features suggest that, despite the simple morphology that can
be directly observed in fossil specimens of Ikaria, this organism was
remarkably complex, compared with contemporaneous Ediacara
Biota taxa.
Molecular phylogenetic analysis of modern metazoans dem-
onstrates that developmental programing is highly conserved
between disparate groups. Initially, this led to hypotheses that
the last common ancestor (LCA) of bilaterians (animals with two
openings and a through-gut) was relatively complex, containing
many of the features common to a variety of such groups, in-
cluding eyes, segmentation, appendages, and a heart (41–43).
Expansion of this analysis to nonbilaterian animals and their
closest single-celled ancestors instead indicates that components
of these conserved developmental pathways have deep ancestry
(see ref. 44 for discussion). Combined with recent evidence for a
sister-group relationship between Xenacoelamorpha and Bilateria,
this suggests that the bilaterian LCA was a simple, small, mobile
organism with anterior/posterior differentiation and limited
sensory abilities (44–49). Remarkably, these predictions agree
closely with the characters identified here for Ikaria.
Evans et al.
 Recognition of the totality of traits in Ikaria is reliant on both
body and associated trace fossils. Given the simple morphology
of Ikaria, it is unlikely that we would be able to confidently assign
it to the Bilateria, or even Metazoa, without this relationship.
This is consistent with hypotheses that the apparent gap between
molecular clock predictions for the early divergence of bilaterians
and their later appearance in the fossil record is the result of their
predicted simple morphology (44, 49). Thus, similar prephylum,
total group bilaterians may be found elsewhere in the Precambrian
fossil record; Ikaria provides a search image for the future iden-
tification of such forms.
The stratigraphic position of Helminthoidichnites suggests that
Ikaria is the oldest total group bilaterian from the fossil record of
South Australia. Ikaria represents a rare example in early animal
evolution where phylogenetic predictions correspond directly
with the fossil record. Further, the global distribution and rec-
ognition of Helminthoidichnites in Cambrian strata (12, 13) is
distinct from the overwhelming majority of the Ediacara Biota.
While Ikaria is not necessarily responsible for the production of
all examples of Helminthoidichnites, it is likely that Ikaria and/or
related taxa are rare fossil animals that existed across the
Ediacaran–Cambrian boundary.
Materials and Methods
Fossil specimens from the National Heritage fossil site at Nilpena remain in
the field due to occurrence on large (square-meter to square-decameter
scale) bedding planes (18). These specimens are identified by bed and field
numbers (e.g., 1T-A 001). Float specimens from Nilpena are collected and
housed at the South Australia Museum in Adelaide and identified by
P numbers.
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Evans et al.
Specimens of Ikaria and Helminthoidichnites were documented through
digital photography, using a Pentax K-50 digital single-lens reflex, and latex
molds. Helminthoidichnites width was measured by using digital calipers
directly on fossil specimens. Detailed morphological investigation was made
possible by 3D laser scans, collected by using the HDI Compact C506 3D laser
scanner. The accuracy of this scan system is reported to 12 μm. Scans were
processed by using the FlexScan3D software. Measurements were conducted
on 3D scans by using the FlexScan3D software. Screenshots of these scans are
presented in Figs. 1 and 2.
We used the Anderson–Darling test to statistically compare the size fre-
quency distributions of Helminthoidichnites and Ikaria using the freely
available PAST software (https://folk.uio.no/ohammer/past/). For this analy-
sis, we compared the average widths of 606 individual Helminthoidichnites
with the maximum widths of 112 Ikaria from Nilpena (Dataset S1). This
analysis produced a statistically significant P value (>0.05) of 0.448, in-
dicating that we cannot reject the null hypothesis that the two samples are
taken from populations with equal distributions.
Data Availability Statement. All data discussed in this paper are available in
Dataset S1.
ACKNOWLEDGMENTS. This work was supported by NASA Exobiology Pro-
gram Grant NNX14AJ86G (to M.L.D.) and NASA Earth and Space Science
Fellowship Program Grant NNXPLANET17F-0124 (to S.D.E.). S.D.E. and M.L.D.
were supported by the NASA Astrobiology Institute under Cooperative
Agreement NNA15BB03A, issued through the Science Mission Directorate.
DEVELOPMENTAL
We thank R. and J. Fargher for access to the National Heritage Nilpena
Ediacara fossil site on their property, acknowledging that this land lies
within the Adnyamathanha Traditional Lands. Fieldwork was facilitated
BIOLOGY
by M. A. Binnie, M. Droser, R. Droser, M. Dzaugis, M. E. Dzaugis, P. Dzaugis,
M. Ellis, C. Hall, E. Hughes, C. Peddie, J. Perry, D. Rice, R. Surprenant, and
L. Tarhan. We thank D. Erwin and J. Irving for helpful discussion regarding
this manuscript. S. Wasif created Fig. 3.
AND PLANETARY SCIENCES
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