National Science Review

RESEARCH ARTICLE 5: 863–869, 2018

doi: 10.1093/nsr/nwy057
Advance access publication 1 June 2018

GEOSCIENCES

Origin of raptorial feeding in juvenile euarthropods

revealed by a Cambrian radiodontan
Jianni Liu1,∗ , Rudy Lerosey-Aubril2 , Michael Steiner3 , Jason A. Dunlop4 ,

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Degan Shu1,∗ and John R. Paterson2
1 Shaanxi Key

Laboratory of Early
Life and Environment,
The Key Laboratory of
Continental Dynamics,
Department of
Geology, Northwest
University, Xi’an
710069, China;
2 Palaeoscience
Research Centre,
School of
Environmental and
Rural Science,
University of New
England, Armidale,
New South Wales
2351, Australia;
3 Institut für
ABSTRACT
The rapid rise of arthropods during the Cambrian quickly established some clades, such as the euarthropod
stem-group called Radiodonta, as the dominant and most diverse predators in marine ecosystems. Recent
discoveries have shown that the size and dietary ecology of radiodontans are far more diverse than
previously thought, but little is known about the feeding habits of juveniles. Here, we document a very small
(∼18-mm-long), near-complete specimen of the radiodontan Lyrarapax unguispinus from the early
Cambrian Chengjiang Biota of China. This specimen is the smallest radiodontan individual known,
representing a juvenile instar. Its adult-like morphology—especially the fully developed spinose frontal
appendages and tetraradial oral cone—indicates that L. unguispinus was a well-equipped predator at an early
developmental stage, similar to modern raptorial euarthropods, such as mantises, mantis shrimps and
arachnids. This evidence, coupled with the basal phylogenetic position of radiodontans, confirms that
raptorial feeding habits in juvenile euarthropods appeared early in the evolutionary history of the group.
Keywords: Lyrarapax, Radiodonta, Arthropoda, Cambrian, predation, feeding ecology

Geologische INTRODUCTION mouth apparatus and the phylogenetic concept of

Wissenschaften, FU the Radiodonta.
Many Cambrian members of the iconic euarthro-
Berlin, Malteserstraße pod stem-group known as radiodontans (Anomalo-
74–100, Haus D, caris and kin) have been viewed as giant apex preda-
12249 Berlin,
RESULTS AND DISCUSSION
tors ever since their true body plan was revealed
Germany and
over 30 years ago [1–3]. However, more recent find- General morphology
4 Museum für

Naturkunde, Leibniz
Institute for Evolution
and Biodiversity
Science,
Invalidenstrasse 43,
D-10115 Berlin,
Germany
∗ Corresponding
authors. E-mails:
eliljn@nwu.edu.cn;
elidgshu@nwu.edu.cn
Received 17 April
2018; Revised 16
May 2018; Accepted
30 May 2018
ings have shown that the body size of some early
Palaeozoic radiodontans can range from 4 cm to
over 200 cm in length [4–6] and that the highly
variable frontal appendage morphologies are sugges-
tive of a range of feeding modes, from shell-crushing
to filter-feeding [6–11]. Lyrarapax from the early
Cambrian Chengjiang Biota of China represents
the smallest radiodontan taxon, with previously re-
ported body sizes ranging from 4 to 8 cm in length
[4,5]. Until now, the frontal appendage morphol-
ogy of the type species, Lyrarapax unguispinus was
only known from a single, incomplete appendage
[4,5], but the preserved details imply a predatory
function. The juvenile specimen of L. unguispinus de-
scribed here not only provides novel information
on the frontal appendages and feeding mode in this
taxon, but sheds new light on the morphology of the
The small specimen (total body length: 18 mm) of
L. unguispinus documented here (Figs 1–3 and Sup-
plementary Fig. 1, available as Supplementary Data
at NSR online), from the lower Cambrian (Series
2, Stage 3) Yu’anshan Member, Chiungchussu For-
mation of Yunnan Province in China, is relatively
complete and shows features not seen in the type
specimens [4,5], including a fully articulated frontal
appendage (FA) and a sclerotized oral cone; see
Supplementary Data at NSR online, for a detailed
description and taxonomic discussion. The FA com-
prises 12 podomeres (P1–12): P2–6 bear complex
endites of alternating sizes, but no dorsal spines;
P7–11 are devoid of endites, but have stout dor-
sal spines; and P12 has a short, robust distal claw
(Fig. 2 and Supplementary Fig. 1a, available as Sup-
plementary Data at NSR online). Of note is the


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Figure 1. Juvenile specimen of Lyrarapax unguispinus. (a) and (b) Part of a near-complete specimen (XDMU-133) and inter-
pretative drawing. (c) and (d) Counterpart (mirrored) and interpretative drawing. Scale bar in (a) is 2 mm and applies to all
images. af, anterior flange; as, anterior sclerite; cr, caudal rami; es, eye stalk; ey, eye; fa, frontal appendage; fg, foregut;
ls, lateral sclerite; mhg, undifferentiated midgut-hindgut; nm, neck muscle; oc, oral cone; sr, strengthening rays; tf, trunk
flap; tm, trunk muscle. Colours: dark grey, cephalic sclerites; dark purple, foregut; green, frontal appendages; light purple,
undifferentiated midgut-hindgut; pink, muscle blocks; yellow, oral cone.

hypertrophied endite on P2 that hosts at least seven
anterior auxiliary spines (Fig. 2c). The oral cone
consists of many circumoral plates, including four
large, perpendicularly arranged plates with surfi-
cial nodes (Fig. 3a and b). This tetraradial arrange-
ment of large, node-bearing plates intercalated be-
tween a series of smaller plates is very similar to
the Peytoia/Hurdia-like oral cone recently described
from the early Cambrian Guanshan Lagerstätte of
China [12].
RESEARCH ARTICLE
Figure 2. Frontal appendage of Lyrarapax unguispinus. (a) and (b) Photographs of part and counterpart. (c) and (d) Back-
scattered electron micrographs of part and counterpart, showing the anterior sclerite (as), auxiliary (arrowheads) and dorsal
(Ds) spines, endites (E), distal claw (dc) and podomere (P) boundaries. All scale bars: 1 mm.
Significance of the oral cone
The presence of a tetraradial oral cone in L. un-
guispinus has important implications for radiodon-
tan systematics. Previously documented specimens
of Lyrarapax from the Chengjiang Biota do not
display a plated mouth apparatus, but rather a se-
ries of concentric ridges and furrows [4,5]. Based
on the occurrence of this feature in one specimen
of each species (L. unguispinus and L. trilobus), a
lack of circumoral plates was interpreted as a mor-
phological characteristic of Lyrarapax, rather than
a taphonomic artefact [5]. This prompted a revised
generic diagnosis, and also challenged the original
concept of the Radiodonta [3,5]. The oral cone
in the new specimen of L. unguispinus (Fig. 3a
and b) indicates that its absence in other speci-
mens of Lyrarapax is indeed preservational. The
displacement or lack of other external, sclerotized
features (e.g. FAs) suggests that the previously illus-
trated Lyrarapax specimens represent carcasses that
have suffered from some preferential decay of ar-
ticulating membrane and post-mortem disturbance
[13–15]; exuvia can be ruled out, as all specimens
preserve internal labile tissues. Thus, an alternative
interpretation of the oral structures previously de-
Liu et al. 865
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scribed for L. unguispinus (Figs 1a–d, f in [4]) and
L. trilobus (Figs 1.3, 1.5, 2.2, 2.3 in [5]) is that
they represent soft tissues—possibly the pharynx
and its musculature—internally located behind the
oral cone; the ‘triangular areas’ seen in the holo-
type (Fig. 1f in [4]) may represent impressions of,
or even attachment points for, the large plates of
the oral cone. Moreover, isolated radiodontan oral
cones have been found in most Cambrian Konservat-
Lagerstätten [9,12,16], demonstrating that they were
often disarticulated from the body during ecdysis
or after death, and later subjected to biostratinomic
sorting.
The discovery of a sclerotized oral cone in
Lyrarapax demonstrates that the presence of ra-
dial circumoral plates remains a consistent trait
of the Radiodonta (particularly as a single coher-
ent unit) and indeed a characteristic feature of
lower stem-group euarthropods [17,18] and other
ecdysozoans [19]. Also, our phylogenetic analysis
of radiodontans based on an updated version of
a recent dataset [6] (available as Supplementary
Data at NSR online) retrieves Lyrarapax nested
within a monophyletic Amplectobeluidae (Supple-
mentary Fig. 2, available as Supplementary Data at
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Figure 3. Mouth apparatus and gut of Lyrarapax unguispinus. (a) and (b) Oral cone showing tetraradial arrangement of large
plates. (a) Detail showing clear impressions of large and small plates. (b) Outlines of preserved plates and nodes. (c) and
(d) Complete gut with impressions of segment boundaries (arrowheads) in the foregut and possible midgut digestive glands
(asterisks); (c) part and (d) counterpart (mirrored). All scale bars: 1 mm. an, anus; fg, foregut; lp, large plate; m, mouth; mhg,
undifferentiated midgut-hindgut; no, node; oc, oral cone; sp, small plate.

NSR online). The tetraradial arrangement appears to
be plesiomorphic for Radiodonta, and the presence
of node-bearing plates is a synapomorphy of Anoma-
locarididae + Amplectobeluidae [12,20].
Raptorial feeding in adult and juvenile
radiodontans
Of the diverse radiodontan FA morphologies
[5,6,8,10], those of amplectobeluids appear the
best suited for grasping and manipulating prey,
characterized by a proximal hypertrophied en-
dite and a series of robust dorsal spines distally
[4,5,8,20–22] (Supplementary Fig. 3, available as
Supplementary Data at NSR online). Lack of an
articulation joint at the base of the hypertrophied
endite indicates that it may have functioned as the
rigid part of a ‘claw’, with the more flexible portion
of the FA represented by the distal podomeres, thus
permitting pincer-like capture of prey [2,8]. The
stout, curved dorsal spines on the distal podomeres,
when curled inwards, would have aided in securing
the prey [8]. Also, the combination of a ‘dorsal kink’
in the proximal portion of the FAs (Fig. 4b and
Supplementary Fig. 3, available as Supplementary
Data at NSR online) and reduced head sclerites may
have afforded the raptorial appendages a greater
range of motion and degree of flexibility for striking
and seizing prey [23]. As described above, the FA of
L. unguispinus shows a pronounced morphological
and therefore functional differentiation along its
proximo–distal axis. The proximal podomeres are
armed with complex endites, the smaller of which
would have converged towards the serrated margin
of the hypertrophied spine during flexion (Fig. 2 and
Supplementary Fig. 1a, available as Supplementary
RESEARCH ARTICLE
Figure 4. Reconstructions of Lyrarapax unguispinus. (a) Complete body in dorsal view. (b) Frontal appendage in lateral view,
showing the spinosity pattern along the proximo–distal axis and dorsal kink (arrow). A more proximal, long podomere is
suggested based on the morphology of Amplectobelua symbrachiata. (c) Artistic representation of the animal.
Data at NSR online). This likely resulted in crushing
or slicing of prey before ingestion; the similar
FAs of Amplectobelua stephenensis may have had a
comparable function [8]. In contrast, the endites of
A. symbrachiata are more simple and slender and,
while the FA of this species would have undoubt-
edly been used to grab prey, mastication was likely
performed separately by gnathobase-like structures
[20]. Amplectobeluid FAs may have been able
to apply considerable force, as evidenced by the
darker pigmentation of their spines in Chengjiang
specimens (e.g. Supplementary Fig. 3a, available as
Supplementary Data at NSR online; Figs 3.2, 3.4
in [5]; Fig. 1A in [20]). This darker colouration is
often indicative of a local thickening of the cuticle,
as recently demonstrated for the gnathobases of
the Cambrian durophagous euarthropod Sidneyia
inexpectans [24]. However, forceful capture may
have caused occasional damage, as seen in A.
symbrachiata FAs where the more delicate (non-
hypertrophied) endites have broken off in some
cases [20,21].
This raptorial feeding mode in Lyrarapax and
Amplectobelua seems to extend to juveniles as well.
Immature specimens of L. unguispinus (Figs 1 and 2)
and A. symbrachiata (Fig. 3A in [2]; Fig. 125 in [25];
extended data Fig. 1b–d in [4]) show that the FAs
have distinct adult-like morphologies. Also, the pres-
ence of large eyes, well-developed body flaps and an
enlarged gut in juveniles [2,25] (Figs 1, 3c, 3d and
Liu et al. 867
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Supplementary Fig. 1c–e, available as Supplemen-
tary Data at NSR online) suggests that Lyrarapax
and Amplectobelua were already highly mobile visual
predators during the early stages of post-embryonic
development. It is possible that similar predatory
modes occur among juveniles of some other ra-
diodontans, such as Anomalocaris, given the compa-
rable set of traits in adults [17,26,27], but this can
only be confirmed by studying juvenile specimens.
Despite the lack of suitable modern analogues
for reaffirming the proposed functional morphology
and feeding ecology of radiodontans [28], there ex-
ist several predatory euarthropod groups that hunt
using enlarged, spinose raptorial appendages. These
include terrestrial clades such as the amblypygids
(whip spiders) [29,30], uropygids (whip scorpions)
[30] and mantodeans (mantises) [31], as well as the
marine stomatopod crustaceans (mantis shrimps)
[32–35] (Supplementary Figs 4 and 5, available as
Supplementary Data at NSR online). Interestingly,
the highly specialized morphology of grasping ap-
pendages develops very early during the ontogeny of
these taxa.
Perhaps the most striking resemblance to ra-
diodontan FAs are the pedipalps of uropygids [30]
(Supplementary Fig. 5, available as Supplementary
Data at NSR online), which provide one of the best
modern analogues for understanding the functional
morphology of the FAs of Lyrarapax throughout
ontogeny. Both adult and juvenile uropygids use
868 Natl Sci Rev, 2018, Vol. 5, No. 6
their large, curved raptorial pedipalps to catch and
hold prey, with the various stout spines on the tar-
sus, tibia and patella acting like prehensile pincers,
while the opposing gnathobases on the proximal
trochanters crush and masticate the victim [30]. The
robust construction of the pedipalps affords uropy-
gids a varied diet [30,36], but juveniles seem to avoid
prey with hard exoskeletons [30]. This may have also
been the case in Lyrarapax and Amplectobelua, al-
though the reinforced spines in immature FAs may
have allowed juveniles to manipulate hard (possi-
bly even biomineralized) food items. So, while the
FAs of amplectobeluid radiodontans may have func-
tioned in a similar way to the pedipalps of uropy-
gids, consumption of the prey was clearly differ-
ent. Uropygids use their chelicerae to tear tissue
from the prey before passing it to the preoral cav-
ity, where it is liquefied by digestive fluids [30]. For
radiodontans, there is still debate as to whether the
oral cone performed an additional masticatory role
or was used to ingest food via suction [16,28]; also,
A. symbrachiata would have used its gnathobase-like
structures to initially process food before passing it
to the mouth opening [20]. However, given the vari-
able morphologies of the oral cone (as corroborated
by FAs), it is likely that the assorted radiodontan
species used this structure in different ways [12,16].
The variety of radiodontan feeding structures
clearly points to these stem-group euarthropods
having played key, often high-tier, trophic roles
within early Palaeozoic food webs, including the
consumption of zooplankton, as well as nektonic
and benthic fauna [6,8,10–12,16,20,37,38]. While
certain taxa such as Anomalocaris [27] can still
be considered giant apex predators of their time
and capable of consuming large prey, the juveniles
of some radiodontans like Lyrarapax (Fig. 4 and
Supplementary Fig. 6, available as Supplementary
Data at NSR online) demonstrate that predation
in the water column was occurring on a variety of
scales during the Cambrian [10,37,39–41]. On the
smaller scale of prey, taxa such as Tamisiocaris were
likely microphagous suspension feeders of plank-
ton (≥0.5 mm in size) [10], whereas juvenile am-
plectobeluids were probably capable of feeding on
very small (<5-mm) benthic and nektonic prey
items, possibly including sclerotized taxa or biomin-
eralized forms such as trilobites, molluscs and bra-
chiopods. The consistent raptorial feeding habits
of both juvenile and adult Lyrarapax contrast with
the example of another early Cambrian euarthro-
pod: the megacheiran Leanchoilia illecebrosa, where
it is thought that juveniles and adults occupied sep-
arate ecological niches due to differing appendage
morphologies [42]. Hence, the predatory lifestyles
of certain radiodontan offspring adds further tier-
RESEARCH ARTICLE
ing complexity to Cambrian marine food webs, and
would have likely placed extra selective pressures
on animal communities, particularly small benthic
and nektonic prey [28,37]. Intense predation occur-
ring on all scales during the early phase of animal
evolution was undoubtedly a critical driver behind
the morphological and ecological innovations aris-
ing throughout the Cambrian [28].
SUPPLEMENTARY DATA
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Supplementary data are available at NSR online.
ACKNOWLEDGEMENTS
Thanks to E. Baker, J. Haug, S. Huber, C. McLean, M. Seiter and
J. Wolff for discussions and supplying images of modern raptorial
euarthropods; J.-B. Caron and D. Huang for images of amplecto-
beluid frontal appendages; L. Yang for taking photos of A. sym-
brachiata specimens; K. Wang for the artistic 3D reconstructions;
T. Brougham for conducting the phylogenetic analysis; and P.
Van Roy and an anonymous referee for their helpful reviews. J.L.
and D.S. designed the research. J.R.P., R.L.A. and J.L. wrote the
text, and all authors discussed and approved the manuscript. J.L.,
R.L.A. and J.R.P. performed light photography, and M.S. acquired
SEM images. J.L. produced the original camera lucida drawings
for Fig. 1 and R.L.A. created the line drawings in Figs 4a and 4b.
R.L.A. prepared all figures.
FUNDING
This research was supported by Project 973 of the Ministry of
Science and Technology of China (2013837100), the Strategic
Priority Research Program of Chinese Academy of Sciences
(XDB26010104), the National Natural Science Foundation of
China (41222014, 41172023, 41621003, 41102012), the 111
Project and the Ministry of Education of China for Chengjiang
Scholars to J.L., and an Australian Research Council Future
Fellowship (FT120100770) to J.R.P.
Conflict of interest statement. None declared.
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 Origin of raptorial feeding in juvenile euarthropods revealed by a

Cambrian radiodontan

Jianni Liu, Rudy Lerosey-Aubril, Michael Steiner, Jason A. Dunlop, Degan Shu & John R.

Paterson

SUPPLEMENTAL TEXT

SYSTEMATIC PALAEONTOLOGY

Material & Methods. The material studied consists of the part and counterpart of an almost complete, juvenile

specimen of Lyrarapax unguispinus, deposited in the Northwest University (NWU) Museum, Xi’an, China

(XDMU-133). Six frontal appendages of Amplectobelua symbrachiata held in the Early Life Institute (NWU)

were also examined for comparison, including the specimen (ELI-EJ1351A) illustrated in Figure S3A. Digital

images of XDMU-133 were acquired using a Leica IC80HD camera mounted on a Leica M125

stereomicroscope (specimen dry; plain light), and a FEI Quanta 650 scanning electron microscope (back-

scattered; environmental mode). In some cases, up to 10 camera images were taken through a range of focal

planes, then stacked and processed in Adobe Photoshop CS6. Specimen ELI-EJ1351A (dry, uncoated) was

photographed using a Canon EOS 5D Mk IV digital camera equipped with a 100 mm macro-lens. Camera

lucida drawings were made using a Leica M125 stereomicroscope, then digitized using Adobe Photoshop CS6.

All figures were prepared using the latter software.

Terminology used here essentially follows Briggs [1] and Cong et al. [2]. Importantly, ‘proximal(-ly)’

and ‘distal(-ly)’ are used in reference to the larger and smaller extremities, respectively, of both the whole

appendage and the endites. To avoid confusion between these two contexts, auxiliary spines occurring on the

margin of an endite facing the proximal end of the appendage are described as ‘posterior’, the ones located on

the margin facing the distal end of the appendage as ‘anterior’ [1]. The term ‘anterior sclerite’ is preferred over

the term ‘dorsal plate’ of some authors (e.g., Cong et al. [3]), for we regard this structure as homologous to the

anterior sclerite of some deuteropods [4]. Abbreviations used: exs., exsagittally; FA, frontal appendage; P1–12,

podomeres 1–12; T1–8, trunk segments 1 to 8; tr., transverse; sag., sagittally; E2–6, endites of podomeres 2 to 6.
Order Radiodonta Collins, 1996 [5]

Emended diagnosis. Stem euarthropods with a non-biomineralized cuticle, more heavily sclerotized in the

frontal appendages, the oral cone, and the anterior and lateral sclerites. Body subdivided into cephalic and

postcephalic regions based on absence or presence of lateral flaps, respectively. Cephalic region unisegmented,

bearing a pair of frontal appendages, a pair of stalked eyes, an anterior sclerite, and an oral cone; frontal

appendages (FA) with at least nine articulated podomeres, typically bearing endites with auxiliary spines; eyes

compound and large, with stalk nearly as thick as the eye itself; anterior sclerite covering the anterior part of

head, and more or less protruding from it anteriorly; ventral mouth surrounded by radially-arranged plates,

connected to one another in a highly-cohesive structure, the oral cone. Postcephalic region metamerized,

composed of 10–14 post-oral segments; each segment bears one or two pairs of triangular or lanceolate,

typically imbricated flaps, associated with groups of parallel-oriented, elongated lanceolate blades; anteriormost

postcephalic segments may be markedly reduced, thus forming a discrete neck region (emended from [3]).

Remarks. The emended diagnosis above is essentially the same as that of Cong et al. [3], except that it reinstates

the presence of the oral cone as a diagnostic feature of the order, as originally proposed by Collins [5], and in

accordance with the name Radiodonta. As discussed in the main text, we consider the absence of this structure

in some specimens of Lyrarapax [3, 6] as the result of postmortem disarticulation, the juvenile specimen

described herein clearly establishing its original presence in this taxon, as in all radiodontans.

The mouth apparatus of radiodontans is reminiscent of the systems of radially-arranged circumoral

plates/teeth characterizing more basal stem-group euarthropods, other panarthropods (e.g. lobopodians), and

cycloneuralians [7–9], which suggests a deep origin of this feature in the evolutionary history of ecdysozoans.

Yet, the radiodontan mouth apparatus differs from these other mouth apparatuses by the coherent assembly of its

elements into a single unit – the oral cone. Indeed, the circumoral plates of radiodontans are almost always

found associated together as an oral cone, even when this structure is found disarticulated from the body as part

of a moult assemblage, or completely isolated following postmortem disarticulation and redistribution [10–13]

(see fig. 8.1 in [14] and fig. 1k in [15] for possible rare exceptions). Even when more than one ring of plates are

present (e.g. in Hurdia), the inner rings made of small plates are always found attached to the main ring of large

plates [16]. These observations attest to a unique structural integrity of this assemblage of plates, which

contrasts with the non-radiodontan taxa mentioned above, including the relatively closely related Pambdelurion,

in which the arrangement of plates was apparently easily disturbed after death [9].
 Cong et al. [2] illustrated the presence of small sclerotized plates, either smooth or tuberculate, in

Amplectobelua symbrachiata, which they interpreted as disarticulated circumoral plates. Such an easily-

disarticulated mouth apparatus would be a significant departure to the condition hitherto described in

radiodontans. Moreover, it is important to note that their reconstruction of the putative mouth apparatus in this

taxon (their fig. 10) is hypothetical (as acknowledged by the authors) – at present, no specimens showing such

plates radially organized as a mouth apparatus have been described. As such, there is no true indication of a

radial organization of these plates. In fact, the tuberculate plates are aligned in series in certain specimens (figs

1, 5, 8 in [2]), sometimes closely associated with similarly aligned gnathobase-like structures, as if forming parts

of one structure (fig. 5 in [2]). Even when observed in more disorganized assemblages of sclerites, the putative

circumoral plates are typically found in close association with the gnathobase-like structures. This observation is

at odds with the interpretation of these plates as circumoral (i.e. cephalic) and therefore anterior to the segments

of the neck region bearing the gnathobase-like structures. On the other hand, the absence of a typical

radiodontan oral cone in any of the specimens of A. symbrachiata illustrated to date is puzzling. Nonetheless,

and pending more definitive evidence regarding the architecture of the A. symbrachiata mouth apparatus, we

propose to consider the highly-cohesive oral cone of radiodontans as a diagnostic feature.

Genus Lyrarapax Cong, Ma, Hou, Edgecombe and Strausfeld, 2014 [6]

Type species. Lyrarapax unguispinus Cong et al., 2014 [6], from the Yu’anshan Member of the Chiungchussu

Formation, Ercai and Mafang sections, Haikou area, eastern Yunnan Province, China.

Other species. Lyrarapax trilobus Cong et al., 2016 [3], from the Yu’anshan Member of the Chiungchussu

Formation, Jianshan section, Haikou area, eastern Yunnan Province, China.

Diagnosis (emended). Radiodontan exhibiting the following unique combination of characters: anterior sclerite

oval, covering anterior part of head only and protruding from it anteriorly; second(?) podomere of frontal

appendage bearing a massive, blade-shaped endite, equipped with at least seven anterior auxiliary spines

increasing in size distally; neck region well-differentiated, composed of reduced anteriormost four postcephalic

segments; first trunk segment conspicuously larger than any succeeding segments.
Lyrarapax unguispinus Cong, Ma, Hou, Edgecombe and Strausfeld, 2014 [6]

New material. Almost complete juvenile specimen XDMU-133, part and counterpart (Figs 1–3, and

Supplementary Fig. S1).

Locality and age. Yu’anshan Member of the Chiungchussu Formation, Ercai section, Haikou area, eastern

Yunnan Province, China; Cambrian Series 2, Stage 3, Eoredlichia-Wutingaspis Zone.

Description of the new specimen. Except for an eye and a FA, the body is complete and about 18 mm in length

(sag.; from anterior margin of anterior sclerite to base of caudal rami). It exhibits a marked subdivision into a

head (or cephalic region), neck, and trunk (post-cephalic region), and is observed from the ventral side (Fig. 1).

Head region subrectangular, slightly rotated clockwise relative to post-cephalic region, and exhibiting an

anterior sclerite, a pair of lateral sclerites, a stalked eye, an oral cone, and a FA. Anterior sclerite oval in outline,

protruding forwards beyond insertion sites of FAs and left eye (Fig. 1A–D). The eye is large and elongate,

subtriangular in outline, prolonged anteromedially by short and robust stalk (Fig. 1C and D). Insertion site of

stalk on head covered by left lateral sclerite, but possibly slightly posterior to insertion site of left FA. Few

polygonal structures, possibly representing lenses, are visible in distalmost part of eye. Paired, lateral sclerites

elongate (exs.) and grossly reniform (Fig. 1A–D). Oral cone covering posterior three-fourths of head region

ventrally, composed of four large, perpendicularly-arranged, node-bearing plates, separated from one another by

smaller plates (Figs 1, 3A and B); the total number and detailed organization of smaller plates is unknown due to

poor preservation. Boundary with neck region close to posterior margin of oral cone.

Right FA comprising 12 visible podomeres (P1–12), assuming that the most proximal portion truly

represents a single, distinct, possibly spineless podomere (Figs 1, 2, and Supplementary Fig. S1A). P2–12

exhibit different spinosity patterns (Fig. 2D): P2–6 bear endites (E2–6), both alternating in size and decreasing

in size distally, but no dorsal spines; P7–11 are devoid of endites, but bear stout dorsal spines; and P12

apparently bears a short and robust distal claw. E2 particularly large, roughly as long as distal portion of FA

(P7–12) and slightly curved toward it; it bears one posterior and at least seven anterior auxiliary spines on its

basal half only, the anterior ones increasing in size distally (Fig. 2C and Supplementary Fig. S1A). E3 barely a

fifth of E2, with a single auxiliary spine on both anterior and posterior margins (Fig. 2D and Supplementary Fig.

S1A). E4 long, about half the length of E2, and bearing a pair of posterior auxiliary spines only. E5 about a tenth

of E2, of unclear morphology, but probably similar to E3 (i.e. with one posterior spine and one anterior auxiliary
spine). E6 roughly equal to a fifth of E2 in length, extending distally no further than the two auxiliary spines

projecting from its posterior margin, and strongly inclined towards distal end of FA.

Neck region composed of at least three, probably four narrow (sag.) segments, as evidenced by faint

ridges marking segment boundaries, and a series of tiny lateral flaps associated with mineralized structures

increasing in size posteriorly (Fig. 1C and D, and Supplementary Fig. S1B). The fibrous texture and position of

these latter structures suggest they might represent remains of metamerically-disposed muscles (see below).

Trunk region semi-circular in outline, comprising eight segments (T1–8) with laterally splayed flaps and

a pair of caudal rami (Fig. 1). Segment boundaries unclear in main trunk, but number of segments inferred from

number of flap pairs and dark-coloured areas at flap insertion sites (Fig. 1A and B). T1 particularly large, about

twice as wide (tr.) as long (exs; measured at flap bases), and bearing long lateral flaps projecting posterolaterally

(ca. 125°). Each flap displays a raised (in ventral view), typically dark-coloured anterior flange, about a fifth of

flap length (exs.; Fig. 1, and Supplementary Fig. S1C–E), which is bordered anteriorly by a flat, narrow rim

proximally and thins out distally. It appears somewhat striated, but also bears more clearly defined linear

impressions (‘strengthening rays’ or ‘transverse lines’ of previous authors; see [3]), sometimes extending to

main part of flap. T1 right flap also subdivided by a marked change in relief into a proximal part and a distal one

(Supplementary Fig. S1E). T2–8 essentially similar to T1 in morphology, but decreasing in length (exs.)—

abruptly between T1 and T2, progressively thereafter—and gently in width (tr.) posteriorly, the semi-circular

outline of trunk region chiefly resulting from rapid reduction of lateral flaps posteriorly. Trunk terminated by

pair of spiniform caudal rami that project posteriorly and to the left (Fig. 1A and B) – there is no clear evidence

of the presence of caudal fans forming a tailfan in this specimen.

Internal trunk structures include pairs of large purple/brown-pigment areas (one pair per post-cephalic

segment; Fig. 1). The latter features are located at mid-length (exs.) of segments abaxially, at insertion sites of

lateral flaps and within proximal parts of the latter. They decrease in size posteriorly, and are aligned with the

mineralized patches occurring in the neck region – accordingly, they are regarded as comparable, though

differently preserved anatomical features. Structures of similar shapes and locations in Anomalocaris [17],

Lyrarapax [3, 6], and Peytoia [12] have been convincingly interpreted as muscles.

Gut tract extends from oral cone anteriorly to posterior tip of main trunk posteriorly, occupying median

half of neck and median third of trunk (Figs. 1, 3C and D, Supplementary Fig. S1B). It is subdivided into two

main parts, interpreted as foregut and undifferentiated midgut-hindgut, by strong constriction close to T2/T3

boundary (Fig. 1). Foregut rapidly widening posterior to mouth to reach maximum width at mid-length (sag.) of
T1, then progressively tapering posteriorly. It is associated with a rather thick layer of iron oxides, exhibiting

more or less continuous transverse lines interpreted as impressions of segment boundaries (Fig. 3C and D, and

Supplementary Fig. S1B; see [17] for comparable features in Anomalocaris). Undifferentiated midgut-hindgut

mostly represented by darker colouration of the rock. In specimen part (Fig. 3C), its general outline suggests an

abrupt widening of the digestive tract posteriorly to reach, at mid-length of T3, a width comparable to maximum

width of foregut; the tract then progressively tapers to T8, where it represents no more than a thick (tr.) pink-

coloured sagittal line. In specimen counterpart (Fig. 3D), this posterior gut region seems to be composed of at

least six pairs of large ovoid structures projecting posteriorly or postero-laterally (large midgut glands?), rather

than representing a continuous gut tract.

Taxonomic assignment. The presence of an oval anterior sclerite in a distinctly anterior position, a well-

differentiated segmented neck region, and a FA bearing a hypertrophied endite (E2) with a denticulate anterior

margin, indicates that the juvenile specimen described above belongs to the genus Lyrarapax. This genus

currently comprises two species, L. unguispinus Cong et al. 2014 and L. trilobus Cong et al. 2016, which

essentially differ from one another by details of the spinosity patterns of their FAs. The fact that the FA

morphologies of the two species are known only from their imperfectly-preserved holotypes, and that these two

specimens clearly differ in size (see figures 3.5 and 3.6 in [3]) calls for some caution in considering the presence

of two distinct Lyrarapax species in the lower Cambrian strata of the Haikou area.

The juvenile specimen described herein greatly resembles the holotype of L. unguispinus [6], the only

Lyrarapax species known from the Ercai section [3]. This is particularly true of its FA, which exhibits rather

high podomere height/length ratios, and endites on P2–6 (= ‘podomeres 3 to 7’ of Cong et al. [3]) that are all

equipped with auxiliary spines – these features supposedly differentiate L. unguispinus from L. trilobus. Despite

its complexity, the FA spinosity pattern described above and that exhibited by the holotype of L. unguispinus are

strikingly similar. The only differences concern the presence or absence of a few auxiliary spines in each

specimen, such as the posterior auxiliary spine of E2 in the juvenile specimen (actually possibly present, but

imperfectly preserved or prepared in the holotype; see figure 3.4 in [3]) or the third posterior auxiliary spine in

E3 and E5 in the holotype specimen. These minor discrepancies are regarded as the result of imperfect

preservation, ontogenetic variation, or perhaps both. Indeed, the FA of the new specimen is particularly small,

being about 20% and 40% smaller than that of the holotypes of L. unguispinus and L. trilobus, respectively (as

estimated from the length of P2–6).

 A diagnostic feature of Lyrarapax is the presence of a four-segmented neck region, but all neck segments

are not always easily differentiated, especially when the flaps of the posteriormost neck segment are concealed

under the large flaps of the anteriormost trunk segment [3]. This fact, or simply the size and preservation of the

new specimen, could easily explain why only three neck segments are recognized with confidence

(Supplementary Fig. S1B). In possessing eight trunk segments, including a noticeably larger anteriormost

segment, the juvenile specimen also strongly resembles other L. unguispinus specimens. According to Cong et

al. [6], the number of trunk segments in this species is “at least eight and probably as many as eleven”, as

deduced from the number of muscle masses observed in the left (eight) and right (eleven) sides of the holotype.

This difference between the two sides of this individual is obviously taphonomic, since it concerns bilaterally

symmetrical structures. This preservation is best understood in light of the recent documentation of limb

musculature in Pambdelurion, an early Cambrian stem-group euarthropod from Sirius Passet in Greenland [18].

In this taxon, each muscle mass is actually composed of two (or more) bundles, which are sometimes preserved

as distinct elements. This is what might occur in the posterior left trunk region of the holotype of L. unguispinus,

which exhibits multiple small muscle masses (discrete bundles) in place of the few large muscle blocks present

in the right region. If correct, the number of main blocks is eight, which likely corresponds to the actual number

of trunk segments in this species. It is noteworthy that some of the pigmented areas on the new specimen also

seem to comprise two distinct elements (darker areas; Fig. 1). In previously described specimens of L.

unguispinus, most, if not all lateral trunk flaps appear to be folded ventrally, roughly at their mid-length (tr.). In

the new specimen, this region of each flap is associated with the abrupt topographical change (Supplementary

Fig. S1E), which explains why the folding occurred in this area. As to the reason for the greater flexibility of the

flaps in previously described specimens (including the holotype), it likely relates to decay-related softening of

the cuticle, which provides further support for the interpretation of these specimens as rather degraded carcasses.

In summary, the new specimen can be confidently regarded as a juvenile individual of L. unguispinus,

and as such provides critical new insights into the anatomy of this taxon and its taphonomy-related

morphological variations. Its FA comprises twelve podomeres organized into two morpho-functional units: a

proximal unit (P2–6) with complex opposing endites forming a particularly efficient crushing/slicing system,

and a distal unit (P7–12) with a smooth ventral margin, but stout dorsal spines and a robust distal claw to assist

with the capture of prey. This elaborate feeding tool is complemented by an oral cone, a complex digestive

system, large eyes, and wide wing-like flexible flaps – all characteristics of a highly mobile, nektobenthic

predator.
PHYLOGENETIC ANALYSIS

Based on the new morphological details provided by the juvenile specimen of Lyrarapax unguispinus, in

addition to recently documented specimens of Amplectobelua symbrachiata [2], we have updated the taxon-

character matrix used by Van Roy et al. [19] (itself derived from that of Vinther et al. [20]) to reassess the

phylogenetic position of these species relative to other radiodontans. The matrix has been provided as a

Supplementary data file in NEXUS format.

Coding. The following characters have been recoded for L. unguispinus and A. symbrachiata using the Van Roy

et al. [19] matrix:

Van Roy et New

Character codings for L. unguispinus
al. [19] coding
8) Mouth: scleral plates overlapping: (0) not overlapping; (1) overlapping in a wheel-
? 1
shaped or pineapple slice array, with enlarged anterior, posterior and lateral plates
9) Mouth: toothplate arrangement: (0) symmetrical, tetraradiate; (1) asymmetrical,
? 0
triradiate
10) Mouth: external surface: (0) smooth; (1) bearing sharp, tooth-like protuberances ? 1
19) Number of appendage pairs (limbs or flap pairs, including frontal appendage): (0) 13
1 0
or fewer; (1) 14 or more
24) Frontal appendage: number of articles^: (0) 14 or more; (1) 13 or fewer ? 1
26) Frontal appendage: length of distal articles: (0) elongate; (1) short and closely packed
? 1
relative to proximal segments
28) Frontal appendage: orientation of terminal spine: (0) terminal spine hooked down;
? 0
(1) terminal spine hooked upward
29) Frontal appendage: terminal segments bearing distally projecting dorsal spines^^: (0)
? 1
absent; (1) present
30) Frontal appendage: form of dorsal spines on terminal segments^^: (0) short and
? 0
robust; (1) long and needle-like
42) Frontal appendage: endites: posterior auxiliary spines: (0) absent; (1) present on one
? 1
or more endites
43) Frontal appendage: endites: auxiliary spines: number of posterior auxiliary spines:
? 1
(0) one (single); (1) two or more
45) Frontal appendage: endites: relative length of lateral and medial spines: (0) subequal;
? 0
(1) medial spine reduced relative to lateral spine
50) Dorsal flap length*: (0) subequal along length of body; (1) anterior flaps about twice
– 1
as long as posterior flaps
53) Transverse rods in flaps: (0) absent; (1) present, transversely oriented ? 1
54) Tail: tailfan: (0) absent; (1) present 1 ?
55) Construction of tailfan: (0) small fluke consisting of a single pair of flaps; (1) large
1 ?
fan, consisting of multiple pairs of flaps
56) Tail: furcae/streamers: (0) absent; (1) present ? 1
58) Lateral carapace elements: (0) absent; (1) present 0 1
60) Extent of central or only carapace element: (0) covering the head; (1) extending in
0 1
front of the head
^ For character 24, we have also recoded Amplectobelua stephenensis as state 1, see [2]
^^ For characters 29 and 30, Caryosyntrips serratus has been recoded as states 1 and 0, respectively, see [21]
* For character 50, we have recoded the following taxa: Opabinia regalis (as state 0), Anomalocaris canadensis (as state 1), Anomalocaris
saron (as state 1).
 Van Roy et
Character codings for A. symbrachiata New coding
al. [19]
4) Eye stalks: (0) absent; (1) present ? 1
7) Mouth: sclerotized oral plates arranged in a radial array: (0) absent; (1) present ? 1
10) Mouth: external surface: (0) smooth; (1) bearing sharp, tooth-like protuberances ? 1
12) Body: division of external surface: (0) undivided; (1) bearing annuli; (2) divided into
? 2
segments
19) Number of appendage pairs (limbs or flap pairs, including frontal appendage): (0) 13
1 0
or fewer; (1) 14 or more
43) Frontal appendage: endites: auxiliary spines: number of posterior auxiliary spines:
1 –
(0) one (single); (1) two or more
50) Dorsal flap length: (0) subequal along length of body; (1) anterior flaps about twice
– 1
as long as posterior flaps
58) Lateral carapace elements: (0) absent; (1) present 0 1
60) Extent of central or only carapace element: (0) covering the head; (1) extending in
0 1
front of the head

Methods. The phylogenetic analysis was performed with TNT v.1.1 [22] using Traditional Branch Swapping

(TBR) under both equal and implied weighting (using k-values of 3, 5, 7 and 9). A separate analysis using

implicit enumeration under equal weighting (as conducted by Van Roy et al. [19]; see their extended data fig.

10) retrieved the same optimal tree length and topology as the TBR analysis.

Results. The analysis under equal weighting produced 1370 most parsimonious trees (MPTs) of tree length 117.

The strict consensus tree (Supplementary Fig. S2A) is 134 steps long and has a consistency index (CI) of 0.545

and retention index (RI) of 0.767, with the topology being similar to that retrieved by Van Roy et al. [19]

(extended data fig. 10). However, our equal weighted analysis using revised character codings (mentioned

above) resulted in the following changes: (1) The node uniting Anomalocarididae + Amplectobeluidae with

Hurdiidae + “Cetiocaridae” collapses, leaving these two clades in an unresolved polytomy with Caryosyntrips

serratus, Cucumericrus decoratus, and other euarthropods (Chengjiangocaris kunmingensis + Olenoides

serratus); (2) The node uniting Hurdia species and Stanleycaris hirpex collapses into a polytomy with the other

hurdiids (with the exception of the more basal taxon referred to as ‘Hurdiid appendage Fezouata’); and (3) A

more resolved Amplectobeluidae, showing Amplectobelua stephenensis and Lyrarapax unguispinus as sister

taxa, which Van Roy et al. [19] retrieved only under implied weights. The collapse of the nodes mentioned in

points 1 and 2 is not surprising, given the poor bootstrap and jackknife values produced by the Van Roy et al.

[19] (extended data fig. 10) analysis. The close relationship between A. stephenensis and L. unguispinus is based

on both taxa sharing a similar number of FA podomeres and a hypertrophied endite with a serrated anterior

margin.
 The implied weights analysis, using a range of k-values (3, 5, 7, 9), resulted in a strict consensus

topology that is identical for all four trees (Supplementary Fig. S2B; tree length 123 steps, CI = 0.593, RI =

0.809). All runs returned 45 MPTs, except for k = 5 that produced 41 MPTs. The topology is much closer to that

resulting from the equal weights analysis conducted by Van Roy et al. [19] (extended data fig. 10). The only

difference with the implied weights tree produced here is the increased resolution within the clade consisting of

Anomalocarididae + Amplectobeluidae (including A. stephenensis and L. unguispinus as sister taxa), and the

more basal position of Paranomalocaris multisegmentalis relative to Anomalocarididae + Amplectobeluidae;

Van Roy et al. [19] also recovered these relationships under implied weights.

The results of the equal and implied weights analyses suggest that a tetraradial oral cone is likely to be

the plesiomorphic condition in radiodontans, given its presence in the Amplectobeluidae (Lyrarapax

unguispinus and possibly Amplectobelua symbrachiata) and Hurdiidae (Hurdia victoria and Peytoia nathorsti),

with the triradial arrangement being a derived feature of the Anomalocarididae. However, this evolutionary

scenario can only be confirmed with further discoveries of articulated specimens or body-part (especially FA

and oral cone) associations across a broader range of taxa. Moreover, oral cones with node-bearing plates seem

to be a unique feature among anomalocaridids and amplectobeluids. The combination of having a tetraradial

arrangement with node-bearing plates is known only in L. unguispinus, possibly A. symbrachiata (although the

circumoral plate arrangement is conjectural; see figure 10 in [2]), and an isolated oral cone from the Guanshan

Lagerstätte [13]. The Guanshan Lagerstätte has yielded two radiodontans, Anomalocaris kunmingensis and

Paranomalocaris multisegmentalis [23]. As noted by Zeng et al. [13], the oral cone they described likely

belongs to A. kunmingensis, for this taxon is a lot more common (>100 specimens) that P. multisegmentalis (a

single specimen). Associating the Guanshan oral cone with either of these species is not at odds with their

phylogenetic placement, even under implied weights where P. multisegmentalis is a basal sister taxon to

Anomalocarididae + Amplectobeluidae (Supplementary Fig. S2B). However, an assignment to A. kunmingensis

appears more likely, considering its phylogenetic position within the Amplectobeluidae.

Despite the variable FA and oral cone morphologies of anomalocaridids and amplectobeluids, these

clades do share other common traits that may represent adaptations for raptorial feeding. This includes the

reduced size of the head elements/sclerites (relative to hurdiids), particularly the oval-shaped anterior sclerite

seen in taxa such as Anomalocaris canadensis [17], Anomalocaris saron [24], Amplectobelua symbrachiata [2],

Lyrarapax unguispinus [6] (Figs 1 and 4 herein), and Lyrarapax trilobus [3]. It has been suggested that the

relatively small head elements/sclerites in anomalocaridids and amplectobeluids permitted the FAs to have a
greater range of motion to aid in prey capture, while also providing some level of protection during an attack

[16, 25]. Interestingly, many members of these clades also exhibit a ‘dorsal kink’ in the proximal portion of their

FAs (Fig. 4B and Supplementary Fig. S3) that may have allowed these appendages to flex backwards to assist

with striking and seizing prey.

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 SUPPLEMENTAL FIGURES

Supplementary Figure 1. Additional morphological details of counterpart. (A) Endites (E2–6) and their auxiliary spines
(arrowheads). (B) Detail of the neck region showing the reduced flaps (nf), their muscles (asterisks), and segment boundaries. (C, D)
Details of trunk flaps showing the anterior flange (af) and strengthening rays (arrowheads). (E) Right side of trunk illustrating the breaks
in slope (arrowheads) between proximal and distal parts of trunk flaps (tf). All scale bars 1 mm.
Supplementary Figure 2. Results of the phylogenetic analysis. (A) Strict consensus of 1370 most parsimonious trees (MPTs) under
equal weighting (consistency index (CI) = 0.545; retention index (RI) = 0.767). Bootstrap (1000 replicates; in bold) and jackknife (1000
replicates, P = 36; in italics) values are indicated above nodes, while Bremer support values are below. (B) Strict consensus of 45 MPTs
under implied weighting (CI = 0.593, RI = 0.809) using concavity value k = 3.
Supplementary Figure 3. Frontal appendages of other amplectobeluid radiodontans. (A) Amplectobelua symbrachiata from the
early Cambrian (Series 2, Stage 3) Chengjiang biota of Yunnan Province, China, specimen ELI-EJ1351A in lateral view (mirrored); note
the darker colouration of largest spines. (B) Amplectobelua stephenensis from the mid-Cambrian (Series 3, Stage 5) Burgess Shale
biota of British Columbia, Canada, holotype (ROM 59492) in lateral view (image: J.B. Caron). (C) ‘Anomalocaris’ kunmingensis from
the early Cambrian (Series 2, Stage 4) Guanshan biota of Yunnan Province, China, holotype (NIGP154566b) in lateral view (mirrored)
(image: D. Huang). Arrows mark position of dorsal kink. All scale bars 5 mm.
Supplementary Figure 4. Juvenile and adult morphologies of modern raptorial euarthropods. (A, B) Whip spider (Chelicerata:
Amblypygi) Phrynichus ceylonicus. (A) Juvenile (deutonymph) (image: C. McLean). (B) Adult female (image: M. Seiter). (C, D) Mantis
shrimps (Crustacea: Stomatopoda). (C) Juvenile (pseudozoea larva; species unknown) (image: J. Haug). (D) Adult specimen of Squilla
rugosa (image: FWC Fish and Wildlife Research Institute, CC-BY-2.0). (E, F) Mantis insect (Hexapoda: Mantodea) Sphodromantis
viridus (images: E. Baker). (E) Juvenile (nymph); Natural History Museum (NHM, London) specimen NHMUK010212122. (F) Adult;
NHM specimen NHMUK010212121. Scale bars, 2 mm (A, E), 10 mm (B), 500 µm (C), 5 mm (D, F).
Supplementary Figure 5. Juvenile and adult morphologies of whip scorpions (Chelicerata: Uropygi). (A) Adult specimen of
Thelyphonus sp. from Malaysia, showing the robust raptorial appendages (pedipalps) (image: B. Dupont, CC-BY-SA-2.0). (B, C)
Typopeltis crucifer from Taiwan (images: S. Huber). (B) Adult female carrying prenymphs. (C) Juvenile (protonymph). ch, chelicerae; fe,
femur; pa, patella; ta, tarsus; ti, tibia; tr, trochanter. Scale bars, 2 mm (A, C), 10 mm (B).
Supplementary Figure 6. Artistic representation of juvenile (foreground) and adult (background) Lyrarapax unguispinus
hunting in the water column. Relative sizes are based on the smallest (length: 1.8 cm) and largest (length: 8 cm) known specimens of
L. unguispinus.