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Haootia quadriformis n. gen., n. sp., interpreted as a muscular cnidarian

impression from the late Ediacaran Period (~560 Ma)

Article  in  Proceedings of the Royal Society B: Biological Sciences · August 2014

DOI: 10.1098/rspb.2014.1202

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Haootia quadriformis n. gen., n. sp., interpreted as a muscular

cnidarian impression from the Late Ediacaran period (approx.
560 Ma)
Alexander G. Liu, Jack J. Matthews, Latha R. Menon, Duncan McIlroy and Martin D. Brasier
Proc. R. Soc. B 2014 281, 20141202, published 27 August 2014

Supplementary data "Data Supplement"

http://rspb.royalsocietypublishing.org/content/suppl/2014/08/26/rspb.2014.1202.DC1.h
tml

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Haootia quadriformis n. gen., n. sp.,

interpreted as a muscular cnidarian
impression from the Late Ediacaran
rspb.royalsocietypublishing.org
period (approx. 560 Ma)
Alexander G. Liu1, Jack J. Matthews2, Latha R. Menon2, Duncan McIlroy3
and Martin D. Brasier2,3
1
Research 2
Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, UK
Department of Earth Sciences, University of Oxford, South Parks Road, Oxford OX1 3AN, UK
3
Department of Earth Sciences, Memorial University of Newfoundland, 300 Prince Philip Drive,
Cite this article: Liu AG, Matthews JJ, Menon
St John’s, Newfoundland and Labrador, Canada A1B 3X5
LR, McIlroy D, Brasier MD. 2014 Haootia
quadriformis n. gen., n. sp., interpreted as a Muscle tissue is a fundamentally eumetazoan attribute. The oldest evidence for
muscular cnidarian impression from the Late fossilized muscular tissue before the Early Cambrian has hitherto remai-
Ediacaran period (approx. 560 Ma). Proc. R. ned moot, being reliant upon indirect evidence in the form of Late Ediacaran
ichnofossils. We here report a candidate muscle-bearing organism, Haootia
Soc. B 281: 20141202.
quadriformis n. gen., n. sp., from approximately 560 Ma strata in Newfound-
http://dx.doi.org/10.1098/rspb.2014.1202
land, Canada. This taxon exhibits sediment moulds of twisted, superimposed
fibrous bundles arranged quadrilaterally, extending into four prominent bifur-
cating corner branches. Haootia is distinct from all previously published
contemporaneous Ediacaran macrofossils in its symmetrically fibrous, rather
Received: 16 May 2014 than frondose, architecture. Its bundled fibres, morphology, and taphonomy
Accepted: 5 August 2014 compare well with the muscle fibres of fossil and extant Cnidaria, particularly
the benthic Staurozoa. Haootia quadriformis thus potentially provides the earliest
body fossil evidence for both metazoan musculature, and for Eumetazoa, in the
geological record.
Subject Areas:
evolution, developmental biology,
palaeontology
1. Introduction
Sediments of Late Ediacaran age (approx. 580–541 Ma) record the fossilized
Keywords:
remains of a diverse global assemblage of soft-bodied macro-organisms. The
Ediacaran, metazoan, Newfoundland,
biological affinities of these Late Ediacaran macrofossils remain the subject of con-
Cnidaria, muscle siderable debate (summarized in [1]). Following their initial discovery, Ediacaran
soft-bodied organisms were commonly assigned to metazoan groups (e.g. [2], or
the classification tables in [3], pp. 240–242). However, the revolution in Ediacaran
thinking brought about by the Vendobiont hypothesis of Seilacher [4] led to recon-
Author for correspondence:
sideration of many of those assignments. Recent years have witnessed a trend
Alexander G. Liu
towards interpreting individual taxa as candidate stem- and crown-group meta-
e-mail: agscl2@cam.ac.uk zoans. Described with varying degrees of confidence, these currently include
potential sponges [5–8], anthozoan, hydrozoan and scyphozoan cnidarians
[9–11], ctenophores [12], placozoans [13], early molluscs ([14]; though see [15])
and even ascidian chordates [16]. These fossils are largely found in successions
of approximately 555–541 Ma, in South China, Brazil, the White Sea region
of Russia, Namibia and the Flinders Ranges of South Australia [17,18]. Further
evidence for the presence of metazoans in the Late Ediacaran period, and indirec-
tly for muscular tissue, comes from simple, putatively bilaterian, surface trace
fossils from the previously mentioned localities [19–21], horizontal surface
traces with crescentic internal divisions made by motile, muscular organisms
Electronic supplementary material is available [22,23] approximately 565 Ma [24], and vertical equilibration traces from
Newfoundland [23]. Prior to 565 Ma, the potential fossil record of animals is
at http://dx.doi.org/10.1098/rspb.2014.1202 or
restricted to claims for biomarkers (e.g. demosponge steranes of more than
via http://rspb.royalsocietypublishing.org.

& 2014 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution
License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original
author and source are credited.
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635 Ma [25]; though see [26]); various specimens interpreted as Peninsula (figure 1f; electronic supplementary material, figures 2
possible sponges from the Early and Middle Neoproterozoic S1 and S5; designated the paratype). The smaller paratype speci-

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([27–29]; though see [8]); and traces of contested age and men has been preserved in lateral view and displays an
origin [30–32]. The absence of clear metazoan body fossils anchoring support structure, lineated stem and a furrowed
until the latest Ediacaran Period renders these earliest reports body with apparent branches (figure 1f; electronic supplemen-
open to debate. Independent estimates for the first appearance tary material, figure S5).
of animals in the Neoproterozoic vary widely, but recent mol- Description. The non-retrodeformed holotype bears a
ecular phylogenetic studies predict that most stem-group discoidal structure 56  37 mm in diameter, preserved in nega-
divergences between extant metazoan phyla occurred within tive epirelief. The disc interior is smooth, apart from faint
the Cryogenian and Ediacaran Periods [33]. concentric ridges at its outer margin (figure 1a), and a small
Newfoundland, in eastern Canada, contains some of the slightly raised central structure of 9 mm diameter with several
oldest non-algal Ediacaran macrofossil assemblages, dated to tight concentric rings (figure 1e). This central structure appears

Proc. R. Soc. B 281: 20141202

approximately 579–560 Ma [34]. Although ichnological evi- to form the attachment point for a short 7-mm wide, lineated
dence for the presence of metazoans in assemblages of this stalk-like structure, 32 mm in length, which extends to the
age has been reported [22,23,35], metazoan body plans have centre of the quadrate body (figure 1a). The body is preserved
yet to be convincingly demonstrated. We here report Haootia as a rectangular sheet 49  72 mm in dimension, characterized
quadriformis n. gen., n. sp. (figure 1) from the lower Fermeuse by well-defined positive epirelief linear ridges (fibres) that are
Formation of the Bonavista Peninsula of Newfoundland 100–600 mm wide and have peaks spaced 200 mm–1 mm
(approx. 560 Ma; electronic supplementary material, figure S1 apart. Individual fibres are finely lineated, exhibiting a struc-
and text S1). This organism exhibits structures wholly consist- ture composed of bundles of parallel strands (figure 1a,b). In
ent with collagenous musculature, in the form of twisted and places, these strands split and then re-join (figure 1b). At the
superimposed fibrous bundles arranged in a quadrilaterally four corners of the body, the fibres converge to form bundles
symmetrical pattern. that progress distally into elongate extensions, here termed
branches (figure 1c). Each of the four corner branches bifurcates
up to three times, and taper towards their distal end, with those
fibres that persist distally decreasing in number after each suc-
2. Systematic Palaeontology cessive branching point (figure 1a,c). Branches were originally
Phylum CNIDARIA Hatschek, 1888 [36] flexible, as demonstrated by 1808 changes in direction of some
Genus HAOOTIA gen. nov. examples to face the predominant flow direction (as inferred
Derivation of name. From the Beothuk (language of the from alignment of nearby unipolar rangeomorphs and Char-
indigenous population of Newfoundland) term Haoot, mean- niodiscus specimens; figure 1a), and by their apparent ability
ing demon, describing the striking appearance of the holotype. to become twisted and rotated (figure 1c). Location of the
Type species. Haootia quadriformis n. gen., n. sp. bulk of the organism down-current of the circular disc
Diagnosis (of genus). Soft-bodied, quadrilaterally sym- in both known specimens is consistent with entrainment by
metrical organism possessing a smooth discoidal structure a flow on the seafloor prior to burial (figure 1a,f; electronic
connected by a relatively short stem to a quadrate body com- supplementary material, figure S5).
prising numerous regularly aligned linear fibres. The fibres Along the margins of the body sheet, between the four
extend laterally across the body, linking adjacent corners. corners, further smaller bundles of linear fibres converge to
Converging fibres extend beyond each corner to form an form small branches that divide dichotomously. Addition-
elongate branch, which divides dichotomously to form smal- ally, along the two shorter edges of the compacted body,
ler, distally tapering sub-branches. Smaller branches also linear fibres running from the adjacent corners combine to
emanate from the lateral margins of the quadrate body, and form bundles that bulge in the middle (figure 1a). By contrast,
these too branch dichotomously. along the two longer edges, the fibres are less obviously clus-
Haootia quadriformis sp. nov. tered into discrete structures, and continue broadly parallel to
Derivation of name. From the Latin quadri (fourfold), and one another.
formis (form), relating to the quadrilateral symmetry of the A prominent linear structure preserved in positive epirelief
organism’s body. runs up the centre-right of the impression, and the fibres of the
Holotype. The original specimen, discovered by M.D.B. in surface of the body appear to drape over it (figure 1a). The
2008, remains uncollected in the field according to provincial narrow morphology of this structure and its similar topo-
law in Newfoundland. A plastotype is held within the collec- graphic relief to the branches leads us to suggest that it
tions of the Oxford University Museum of Natural History, reflects a primary branch from the lower right corner (as seen
specimen OUM ÁT.424/p. in figure 1a), folded beneath the body at the time of burial.
Horizon and locality. From the lower part of the Late Discussion. Haootia quadriformis displays several unique
Ediacaran Fermeuse Formation, St John’s Group [37]. The morphological traits, the most striking of which is an apparently
specimen resides within a turbiditic marine succession (elec- symmetrical, fibrous body with regularly arranged branches
tronic supplementary material, text S1 and figure S2) on the (figure 2b). The superficial impression of bilateral symmetry in
north shore of Back Cove, roughly 1.8 km NNW of the the holotype (figure 2c) was arguably brought about by oblique
town of Melrose, Bonavista Peninsula, Newfoundland, collapse and differential contraction of the body. Biostratinomic
Canada (electronic supplementary material, figure S1). distortion is further enhanced by tectonic stretching. We thus
Diagnosis. As per the genus. infer that the original body was quadrilaterally symmetrical in
Remarks. Haootia quadriformis n. gen., n. sp. is known from the life (figures 2d and 3b), and we suggest that the bedding plane
holotype specimen, and one additional incomplete specimen relationships of the holotype specimen indicate composite pres-
from the Trepassey Formation of Burnt Point, Bonavista ervation of a mould of the base of the anchoring adhesive disc,
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(b) 3
(a)

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Proc. R. Soc. B 281: 20141202
(c)

(d ) (e) (f)

Figure 1. Haootia quadriformis n. gen., n. sp., lower Fermeuse Formation of Back Cove, Bonavista Peninsula, Newfoundland. (a) Haootia quadriformis holotype
specimen. Note the negative-relief central disc, interpreted as a holdfast, and the broadly bilaterally symmetrical bundles of linear ridges, extending into discrete
bifurcating branches. Inferred current direction indicated by the arrow. (b) Fibres running along the right-hand margin of Haootia; each fibre is composed of finer,
thinner fibres. (c) Bottom left corner of Haootia, detailing the connection between a primary bifurcating branch and the main body. Note the twisted fibres along
the branch. (d ) Pinching, bundling and superposition of fibres at the base of a subsidiary branch. (e) The small circular depression at the centre of the disc, showing
mantling parallel fibres forming the base of a short stalk that connects the disc to the body. (f ) Incomplete paratype specimen of H. quadriformis, from the
Trepassey Formation of Burnt Point, Bonavista Peninsula. This specimen is preserved on its side, but clearly displays fibres extending up its stem and around
the body. A small partially buried holdfast disc is arrowed. Scales bars (a,c,f ), 10 mm; (b,d,e), 5 mm.

and the upper surface and internal structure of the body. The the position of the disc upstream of the quadrate body, we
apparent draping of the quadrate body over the disc edge infer that the disc was a tethering structure similar to those of
implies that the body lay above both the disc and stem on the associated frondose taxa (e.g. electronic supplementary
seafloor at the time of burial (figure 1a). On the basis of material, figure S3a–c), and that Haootia was epibenthic.
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(a) (b) 4

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f

c
a

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b
d

(c) (d)

Figure 2. Digitized images of H. quadriformis n. gen., n. sp., emphasizing the convergence of fibrous linear features at the corners of the body, and the symmetry of
the fossil. (a) Photograph of the holotype as it appears in situ. (b) Interpretive sketch of the non-retrodeformed specimen. Labels indicate: (a) muscle bundles,
(b) expanded bundles, (c) ‘contracted’ bundles, (d ) twisting fibres, (e) superimposed fibres and (f ) disc. (c) Digitized overlay of the fossil. Symmetrical regions of
the organism are colour coded. Note the thick bulging of fibres (indicating muscle contraction?) along short axes of the sheet (light green). (d) As in (b), but the
image has been corrected to account for tectonic deformation on the surface by compressing the disc into a perfectly circular structure (cf. [38], though see [39]).
Scale bar, 10 mm.

The complex structure of H. quadriformis, with prominent branching elements [41,42], such features are lacking in Haootia.
bundles of fibres showing consistent directional changes Primocandelabrum sp. [37] (electronic supplementary material,
within a discrete sheet-like structure, is not readily explained figure S6d), a superficially similar contemporaneous rangeo-
by tectonic or sedimentological processes. Unusual environ- morph bearing multiple branches attached by a stem to a disc,
mental taphonomic conditions can also be ruled out, because can be distinguished by its lack of quadrilateral symmetry,
neighbouring specimens of recognizable macrofossil taxa on and its rangeomorph branching. Furthermore, in rare specimens
the bedding planes (e.g. figure 1a) do not differ in preservation where longitudinal ridges are preserved along the length of a
or appearance from those found abundantly throughout the Primocandelabrum [43], such ridges are wider, more broadly
region. All other fossil impressions on these surfaces (electronic spaced and less regular in arrangement than those seen in
supplementary material, figure S3) lack fibrous structures of Haootia. The disc in the holotype Haootia specimen also differs
the kind described here. distinctly from others found on the same surface, being
smoother, with lower topographic relief (figure 1a) and fewer
concentric rings (electronic supplementary material, figure S3).
Examples of putative tissue differentiation in Ediacaran
3. Is this a known Ediacaran macrofossil taxon? macrofossils have typically proved controversial. Structures
Whereas typical frondose Ediacaran taxa possess either leaf-like interpreted as external sheaths and membranes have been
morphologies or some evidence for alternating rangeomorph described in Pteridinium and Rangea from Namibia [44,45],
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‘crumpled’ margins of Karakhtia from the White Sea [52], but 5

(a)
the folds in Karakhtia are irregular in shape and direction, radi-

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ate from the centre of the organism to the outer margin, and
become more finely spaced towards the specimen edges.
Differences are also apparent when considering linear features
associated with ‘mop’ structures in Australia. ‘Mop’ plausibly
results when a disc, embedded in a microbial mat, has been
dragged by unidirectional currents [53] to produce uni-
directional or evenly radiating marks. By contrast, Haootia
fibres form bands that are multidirectional, often running paral-
lel to the margins of the impression and appearing to converge
with neighbouring fibres (figure 1a). Longitudinal furrows are

Proc. R. Soc. B 281: 20141202

known within ribbon-like Harlaniella [54]. Such linear features
demonstrate how individual Ediacaran taxa can exhibit a
variety of putative internal morphologies as a result of differen-
tial taphonomic processes. Such features will also require
explanation, but on the available evidence, we do not consider
Haootia to represent a taphonomic variant of any currently
known Ediacaran taxon. Contemporaneous microbial fabrics
can exhibit linear striated morphologies (e.g. Arumberia [55]),
(b) but are not typically localized in their occurrence, do not pos-
sess a sharp boundary to the impression, and are not known
to form symmetrically arranged bifurcating structures.

4. Metazoan affinities?
Haootia’s size and complex, regular morphology demand con-
sideration of metazoan affinities. Its symmetry and the lack of
evidence for pores or spicules argue against Porifera (following
[8]). The presence of numerous branches, absence of comb rows
and inferred benthic mode of life likewise make comparison
with Ctenophora problematic. Possession of quadrilateral
structure, a central radial disc and fibrous soft tissues, clearly
invite comparison with living and fossil Cnidaria.
Although the extant phylum Cnidaria includes morphologi-
cally and genetically disparate taxa [56,57], their molecular
phylogeny confirms a basal position within the Eumetazoa
[58]. Cnidarians are classically united by the possession of
Figure 3. (a) The extant staurozoan Lucernaria quadricornis, exhibiting a cnidocytes, diploblastic construction and radial symmetry, but
body plan similar to that hypothesized for H. quadriformis n. gen., n. sp. suggestions of a wider variety of symmetry states (e.g. [59–
The Staurozoa are known from a range of marine depositional environments 61]) are supported by genetic arguments for the presence of
and water depths [40]. (b) Artistic reconstruction of H. quadriformis. Scale bilateral symmetry in the eumetazoan common ancestor [62],
bars, 10 mm. and the presence of a mesoderm-like layer has been recognized
in some cnidarian taxa (cf. [63]; electronic supplementary
material, text S2).
and in rare rangeomorphs from Newfoundland [46], although The bundles of fibrous ridges within the body of Haootia
the latter examples likely have a sedimentological origin [47]. compare favourably in size, order and arrangement to the
Such claimed sheaths are typically smooth and lack the fibrous preserved muscular tissue of modern cnidarians. Cnidarians
character of Haootia. The internal anatomy of other Ediacaran can possess smooth and/or striated muscular tissue [63,64]
macrofossils is largely inferred from composite impressions (electronic supplementary material, text S2), both of which
explained by biostratinomic collapse of tissues (e.g. [48], can form fibrous bundles arranged in a similar manner to
fig. 2), or from three-dimensional specimens in-filled by sedi- those in Haootia [65] (figure 3a; electronic supplementary
ment (e.g. [49,50]). However, such typically lobate structures material, figure S6). Rare fossil examples of cnidarian muscu-
do not exhibit the wavy fibrous symmetry of H. quadriformis. lar tissue (e.g. [66 –68]) typically comprise impressions of
Whereas the linear fibrous construction of the alga Flabellophy- regularly arranged ridges (e.g. [67], p. 63, fig. 55). These are
ton from South China and Australia [51] shows some similarity best known in fossil scyphozoan medusae, where coronal
with fibres of Haootia, those fossils lack a large holdfast, a stem- and radial muscles of the sub-umbrella are often grouped
mounted body or quadrilateral symmetry. It could be argued into bundles (e.g. [69]) and are preserved as casts and
that the linear fibres in Haootia result from the deformation or moulds in a taphonomic style similar to that seen in the
twisting of a non-muscular integument, but that cannot explain Ediacaran siliciclastic settings of Newfoundland [70]. The mor-
their presence across the whole body, their multi-directionality phology of soft-bodied fossil cnidarians is typically influenced
or their symmetry. Rough comparison may be made with the by muscle contraction at the time of burial [67]. Twisting and
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overlapping of fossil medusa tentacles [71] also compare clo- food, as with the tentacles of modern cnidarian polyps. 6
sely with Haootia’s flexible branches. Phalloidin fluorescence We see neither a distinct mouth-like structure nor a gastro-

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reveals that the 1–2.5 mm-width smooth muscle fibres in the vascular cavity, so their presence must be inferred at the
extant parasitic hydrozoan Polypodium hydriforme run longitud- centre of the quadrilateral body. Similarly, structures similar
inally up the length of the tentacles [65] in an arrangement to canals or mesenteries are not clearly distinguishable.
strikingly similar to individual fibres in H. quadriformis. Fur- Interpretation of the disc as a benthic holdfast then implies a
thermore, the junction between muscles in the tentacles and polyp-like organism, with a gross body-plan most similar to
those in the body of P. hydriforme produces a similar ‘truncated’ that of living staurozoans (e.g. figure 3). The fibres within
surface to the ridges observed in Haootia (figure 1d; [65], fig. 4a), Haootia are consistent with the positioning of muscular fibres
and individual fibres can also split and/or join one another. in the calyx of modern Staurozoa [85] (figure 3a), being longi-
These morphological and structural similarities lead us to the tudinal within the stalk and branches of the specimen but
conclusion that the fibrous structures preserved within Haootia mainly positioned laterally (i.e. parallel to the margins in a

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may well represent the soft tissue impressions of cnidarian manner analogous to coronal musculature in modern forms
musculature. If so, this specimen significantly pre-dates pre- [84]) in the body. However, the additional marginal branches
viously documented preserved muscular tissues, the oldest in Haootia are unlike anything seen in staurozoans, which typi-
of which are Early Cambrian in age [72,73]. cally possess only eight arms. Haootia also lacks fossilized
Striated muscle fibres have been demonstrated to be pre- evidence for morphological features such as anchors, gonads,
sent in the cubozoan Tripedalia cystophora ([74], fig. 5), and nematocyst clusters or characteristic tissue structures observed
although individual fibres are of smaller magnitude than in histological sections through modern Staurozoa (e.g. ref.
those seen in H. quadriformis, they are nevertheless very simi- [84]). As Haootia is also considerably larger than most extant
lar in gross morphology. Smooth muscle has also been Staurozoa and possesses an unusually large holdfast disc, we
observed to form macroscopic fibrous bundles within the ten- are not in a position to assign it to the class Staurozoa on the
tacles of several scyphozoans [63] and cubozoans [74,75]. basis of available evidence. Cubozoans can also possess bifur-
Distinguishing between bundles of smooth and striated cating tentacles and fourfold symmetry, but extant forms are
muscle cells in the fossil record is not likely to be possible pelagic, not benthic as inferred for Haootia.
when only soft tissue impressions are available for study. Interestingly, symplesiomorphies within the Medusozoa
In the living actinian Metridium, the better-developed have been proposed to include the presence of four intra-
(smooth) longitudinal muscles are notably found in the ecto- mesogleal muscles [40]. The Medusozoa are usually
derm of the tentacles, with circular muscles located in the considered to have a long evolutionary history, with divergence
endoderm ([76], p. 79; contra [77]). This differentiation of from the Octocorallia conservatively estimated to have taken
muscle groups within different tissues may explain why we place at least approximately 571 Ma [86]. If correct, medu-
only see longitudinal ridges along the branches of Haootia, sozoan ancestors, and indeed diverse cnidarian ancestors,
with no clear evidence for circular bands. would be expected within Late Ediacaran marine environ-
The preservation of muscular tissue in the Phanerozoic ments. The suggestion that Staurozoa is the sister group to all
is uncommon and is typically restricted to Konservat other medusozoan classes ([40,87], though see [58]) potentially
Lagerstätten [78]. In many cases, particularly involving arthro- indicates a similarly ancient evolutionary history for that clade.
pod and vertebrate muscle, preservation takes place via Further comparisons with the body plans of extant cnidarians
authigenic replacement of muscular tissues by calcium phos- are limited by our poor understanding of deep sea forms [88],
phate or clay minerals [79], or via sulfurization of organic and the absence of many extinct forms (cf. [59]). Until further
matter [68]. In the Ediacaran, taphonomic processes were morphological evidence is obtained, we therefore suggest
significantly different, and soft tissue preservation was com- that the muscular H. quadriformis n. gen., n. sp. occupied
monly facilitated by the early diagenetic, microbially induced a position within the Cnidaria, and potentially within the
casting of fossil exteriors in framboidal pyrite [47,80] or by stem-group Medusozoa.
rapid burial beneath volcanic ash [81]. Such mouldic preserva-
tion is unusual in the Phanerozoic, but has been documented to
preserve cnidarians (and significantly impressions of their 5. The significance of a cnidarian at
muscular tissue) at several localities [71].
An important consideration is explaining how internal approximately 560 Ma
muscle tissues are preserved in this manner, when in other Interpretation of H. quadriformis as a muscular cnidarian leads
Ediacaran macrofossils we typically only see external mor- us to examine the early fossil record of the phylum Cnidaria.
phology. In taphonomic experiments involving modern Cnidarians appear to have diversified into several major
hydrozoans and scyphozoans, impressions of muscular tissues clades by the Middle Cambrian, as evidenced by the presence
were not preserved [82,83]. However, the absence of microbial of probable anthozoan actinians [89–92] and corals [93–96],
mats on the experimental surfaces [82], and the desiccation of scyphozoans [97], possible hydrozoans and cubozoans [66,98]
specimens [83], precludes direct comparison between those and cnidarians of unknown affinity [99] in Lower and Middle
studies and Ediacaran taphonomic conditions. We suggest Cambrian strata, with conulariids [100] and mass strandings
that rapid degradation of an external integument in Haootia of medusae [101,102] additionally reported in the Upper
(such as the epidermis, less than 50 mm thick in some modern Cambrian (see also [71]). Some of the earliest interpretations
cnidarians [84]) upon death and burial exposed the relatively of the original Ediacara biota of Australia proposed cnidarian
more robust muscular tissues and permitted them to be cast in medusoid affinities for discoidal specimens [103–105], but
the same manner as contemporaneous Ediacaran macrofossils. many of these have since been disputed (e.g. [71,106]).
We infer that the muscle-like fibres seen in Haootia likely Similarly, interpretation of Inaria as an actinian-grade, muscle-
facilitated extension and retraction of branches for gathering bearing polyp [107] has been questioned following taphonomic
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and morphodynamic analysis [77]. Other reports of cnidarians tissue differentiation, feeding strategy, food sources and the 7
in latest Ediacaran rocks include Pambikalbae as a ?hydrozoan complexity of Late Ediacaran ecosystems. Our interpretation

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[108]; interpretation of the tubular fossils Corumbella and of H. quadriformis as a muscular metazoan of cnidarian grade
Vendoconularia as scyphozoans similar to the conulariids arguably represents the earliest known evidence for preser-
[9,11,109]; discussion of the biomineralized genera Cloudina vation of muscular tissue in the geological record, and one of
and Namacalathus as ‘cnidariomorphs’ [110]; and the possible the earliest claims for a eumetazoan (see also [10,114]). Haootia
calcified cnidarian Namapoikia [111]. Fossils from the Late therefore delivers a key calibration point for studies of early
Ediacaran Doushantuo Formation have been tentatively com- Eumetazoan evolution and body symmetry.
pared to tabulates [112,113] and hydrozoans [10]. Elsewhere,
the recent reinterpretation of certain Middle Ediacaran carbon- Acknowledgement. Fieldwork was conducted under permits issued by the
aceous fossils from the Lantian Biota as potential conulariids Department of Tourism, Culture and Recreation, Government of New-
[114] is of interest. Traces of actinian-like locomotion in deep foundland and Labrador. C. Kenchington, P. Wilby, J. Hoyal-Cuthill

Proc. R. Soc. B 281: 20141202

marine sediments approximately 565–560 Ma are also germane and S. Conway-Morris provided helpful discussion regarding compara-
tive material. P. Sargent of the Ocean Sciences Center, MUN, is thanked
here [22,23]. All claims for Neoproterozoic metazoans should
for providing modern comparative material. S. McMahon took high-
be critically assessed on a case-by-case basis, much as with quality images of casts of the specimen, and R. Hooper (MUN) helped
the early sponge fossil record [8]. At the time of writing, how- to identify extant taxa. The manuscript has benefitted from the reviews
ever, the studies cited above clearly indicate morphological of two anonymous reviewers. Plastotype OUM ÁT.424/p is housed at
diversity of fossil cnidarian candidates in the Late Ediacaran/ the Oxford University Museum of Natural History, UK.
Early Cambrian. Such fossils have also been used to help Funding statement. This work was supported by the NSERC (to D.M.);
Natural Environment Research Council (grant no. NE/F008406/1 to
calibrate recent molecular estimates of bilaterian–cnidarian
A.G.L., and NE/J5000045/1 to J.J.M.); a Burdett Coutts grant to
divergence during the Ediacaran Period [33]. J.J.M.; the Cambridge Philosophical Society (a Junior Research Fellow-
Cnidarian-like body fossils from Newfoundland at ship to A.G.L.) and the National Geographic Global Exploration Fund
approximately 560 Ma also raise important questions about (grant no. GEFNE22–11 to A.G.L.).

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