LETTER

doi:10.1038/nature10097
Modern optics in exceptionally preserved eyes of
Early Cambrian arthropods from Australia
Michael S. Y. Lee
1,2
, James B. Jago
1,3
, Diego C. Garc a-Bellido
4
, Gregory D. Edgecombe
5
, James G. Gehling
1
& John R. Paterson
6
Despite the status of the eye as an organ of extreme perfection
1
,
theory suggests that complex eyes canevolve very rapidly
2
. The fossil
record has, until now, been inadequate in providing insight into the
early evolution of eyes during the initial radiation of many animal
groups known as the Cambrian explosion. This is surprising
because Cambrian Burgess-Shale-type deposits are replete with
exquisitely preserved animals, especially arthropods, that possess
eyes
35
. However, with the exception of biomineralized trilobite
eyes, virtually nothing is known about the details of their optical
design. Here we report exceptionally preserved fossil eyes from the
Early Cambrian ( 515 million years ago) Emu Bay Shale of South
Australia, revealing that some of the earliest arthropods possessed
highly advanced compound eyes, each with over 3,000 large omma-
tidial lenses anda specializedbright zone. These are the oldest non-
biomineralized eyes known in such detail, with preservation quality
exceeding that found in the Burgess Shale and Chengjiang deposits.
Non-biomineralized eyes of similar complexity are otherwise
unknown until about 85 million years later
6,7
. The arrangement
and size of the lenses indicate that these eyes belonged to an active
predator that was capable of seeing in low light. The eyes are more
complex than those known from contemporaneous trilobites and
are as advanced as those of many living forms. They provide further
evidence that the Cambrian explosion involved rapid innovation in
fine-scale anatomy as well as gross morphology, and are consistent
with the concept that the development of advanced vision helped to
drive this great evolutionary event
8
.
The anatomy of many Early and Middle Cambrian (,520500
million years (Myr) ago) organisms is known in considerable detail
fromnumerous Burgess-Shale-type deposits worldwide
35
, but the fine
structure of these organisms eyes is poorly known. Details of the visual
surface are known only from certain Cambrian trilobites and some
tiny Orsten arthropods
9
: all of these have compound eyes that are
small inabsolute size and contain fewvisual units (ommatidia) bearing
lenses of uniform size. Yet the evolution of powerful vision has been
proposed as a trigger
8,10
for the Cambrian explosion of animals
3,5
. We
have recovered exceptionally preserved, large compound eyes (Fig. 1)
from the Early Cambrian (Series 2, Stage 4; ,515 Myr ago) Emu Bay
Shale Konservat-Lagerstatte at Buck Quarry, Big Gully, on Kangaroo
Islandin South Australia
1113
. The newfossils represent the oldest non-
trilobite arthropod eyes that show fine detail of the visual surfaces.
All eyes were found isolated (disarticulated) but they are closely com-
parable inmorphology and size, which is consistent withtheir referral to
a single taxon of large arthropods (see below). All specimens bear South
Australian Museum Palaeontology (SAM P) numbers. The visual sur-
faces of three eyes (Fig. 1ac and Supplementary Fig. 1) are relatively
complete and form the basis for the descriptions; the remaining four
(Supplementary Fig. 2) are similar but do not provide extra information.
The fossils are preserved in finely laminated grey mudstone. Scanning
electron microscopy with energy dispersive spectrometry (SEM-EDS)
analysis of the optical surface of specimenP43687 (Supplementary Fig. 3)
detected elevated levels of calcium and phosphorus relative to the
matrix. This indicates that phosphatization of the primarily chitinous
1
South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia.
2
School of Earth and Environmental Sciences, University of Adelaide, South Australia 5005, Australia.
3
School of
Natural and Built Environments, University of South Australia, Mawson Lakes, South Australia 5095, Australia.
4
Departamento de Paleontolog a, Instituto de Geolog a Econo mica/Instituto de Geociencias
(CSIC-UCM), Jose Antonio Novais 2, Madrid 28040, Spain.
5
Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK.
6
Division of Earth Sciences, School of
Environmental and Rural Science, University of New England, Armidale, New South Wales 2351, Australia.
d
0.5 mm
c
p
p
1 mm
1 mm
1 mm
a
p
b
b
b
b
b
Figure 1 | Complex arthropod eyes from the Early Cambrian. ad, Three
fossils of compoundeyes froma large arthropodfromthe Emu Bay Shale, South
Australia (ac), shown in similar hypothesized orientation to the compound
eye of a living predatory arthropod, the robberfly Laphria rufifemorata
(d; anterior view of head). All fossil eyes have large central ommatidial lenses
forming a light-sensitive bright zone, b, and a sclerotized pedestal, p. Because
the fossil eyes are largely symmetrical about the horizontal axis, it is not possible
to determine dorsal and ventral surfaces, and thus whether the eyes are left or
right. All fossils are oriented as if they are left eyes (medial is to the left of the
figure). In b there is a radial tear (white line) with the top portion of the eye
displaced downwards to overlie the main part; extensive wrinkling causes some
central lenses (arrow) to be preserved almost perpendicular to the bedding
plane. South Australian Museum numbers are: a, P43629a; b, P43687;
c, P43658; d, A29-000885.
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cuticle has occurred in some instances, as in the Orsten biota, the only
other Cambrian deposit where non-trilobite eyes are exceptionally pre-
served
9
. Phosphatizationof organic material is relativelycommoninthe
EmuBay Shale, for example, inMyoscolex
14
andIsoxys
11
. Wrinkling and
tearing of the visual surface (for example, Fig. 1b) provide further
evidence that the cuticle was originally non-biomineralized. Despite
flattening of the overall visual surface, finer features such as ommatidial
lenses are preservedinsuperb detail and three-dimensional relief (Fig. 2
and Supplementary Fig. 1).
Each eye comprises a smoothly curved, ovoid visual surface with a
long axis diameter of about 79 mm and a thickly sclerotized pedestal
that is not fully preserved on any specimen. The three most complete
eyes are shown in Fig. 1, in the same presumed orientation as similar
living analogues (for example, the robberfly Laphria and krill
Nematoscelis). The lateral edge of the eye is a smooth arc; the medial
edge, which meets the pedestal, seems to be broadly V-shaped. Each of
the three eyes is preserved in a similar orientation, with the largest
lenses near to the centre and smaller lenses on the margin; this suggests
that each eye was already flattened in this plane during life. The visual
surface contains a dense and highly regular hexagonal array of over
3,000 large lenses. The two best-preserved specimens (P43629a and
P43687, Fig. 1a, b) share many similarities. The eye surface comprises a
dense purplish-black (phosphatized) cuticular layer and is convex at
the margins, indicating that the external surface of the eye cuticle is
exposed. The corneal or lens surfaces of the ommatidia are darker than
the intervening spaces. The lenses are recessed in the central region,
but some near to the margin are elevated; this variation is probably a
taphonomic artefact. The counterpart of P43629a (P43629b) shows a
mould of the presumed external surface in less detail: the lenses are
elevated, there is no dark phosphatizedlayer andthe margins of the eye
are concave. Specimen P43658 (Fig. 1c) is a similar surface to P43629b:
the lenses are likewise elevated, there is no dark phosphatized layer and
the margins are concave.
The lenses of the ommatidia exhibit a similar size gradient across all
specimens. The largest lenses, withabout 150 mmelevational diameter
15
,
are concentrated in the region nearest the pedestal, with size decreasing
to half this diameter (,80 mm) towards the margins (Fig. 2). Lens size
is further reduced to about 60 mm at the extreme margins but com-
pression and oblique orientation make accurate measurement in this
area difficult. The size gradient is not an artefact of compression of
marginal lenses for the following reasons: (1) the measurements given
here refer to the longest diameter across each lenslateral compres-
sionof marginal lenses wouldturna circle intoa progressively narrower
oval but it should not reduce the longer diameter; (2) the lenses get
progressively smaller only a short distance from the centre, well before
any major change in orientation has occurred; and (3) there are deep
wrinkles across P43687, causing some central lenses to be preserved
almost perpendicular to the bedding plane (Fig. 1b, arrow), yet the
longer diameter of these lenses is not reduced. Less pronounced
wrinkles occur in other specimens and similarly do not affect the
gradient of lens sizes. Interommatidial angles cannot be determined
accurately owing tocompression; however, the lenses of the ommatidia
are preserved horizontally nearest to the pedestal but more obliquely
close to the margins. This change in orientation happens abruptly, not
gradually, indicating that the visual surface was relatively flat near to
the pedestal, withcurvature increasing more rapidly near tothe margin
(that is, the visual surface was not uniformly curved).
The Emu Bay Shale eyes are more elaborate than any known
Cambrian visual organ, although comparisons are largely restricted to
the calcitic (and thus optically unusual
8
) eyes of trilobites. The lenses are
not only very numerous andlarge, they are also hexagonally arranged in
a highly regular six-neighbour arrangement: the densest and most effi-
cient packing pattern. In contrast, Early Cambrian eodiscid trilobites
such as Shizhudiscus and Neocobboldia have fewer than 100 lenses,
which are also much smaller (,50mm in diameter) and less regularly
arranged in a less efficient square-grid array
16
. The extremely regular
arrangement of lenses seen here exceeds even that in certain modern
taxa, such as the horseshoe crab Limulus, in which up to one-third of
lenses deviate from hexagonal packing
17
. Eyes with more than 3,000
ommatidia and lenses larger than 60 mm in diameter are otherwise first
known from the biomineralized visual organs of early Ordovician
trilobites, more than 40 million years later (Fig. 3).
The arrangement and size gradient of lenses creates a distinct bright
zone (also called the acute zone or fovea), where the visual field is
sampled with higher light sensitivity (due to large ommatidia) and
possibly a higher acuity (due to what seems to be a more parallel
orientation of ommatidia). Such visual specializations, characteristic
of many modern taxa, are otherwise unknown in the Early Cambrian.
The ratio of lens diameters in the bright zone to lens diameters in the
margin (,2.5:1) exceeds that found in other Cambrian arthropods
(trilobites and cambropachycopids) and is comparable to that in many
modern taxa such as dragonflies, which have ratios of 1.612.71:1
b
c
d
f
g
1
2
3
4
1
2
3
4
Distance along transect (m) Distance along transect (m)
H
e
i
g
h
t

(

m
)
H
e
i
g
h
t

(

m
)
100 200 300 400 500
20
30
1 2 3 4
0
13
100 200 300 400
4.5
0
0
1 2 3 4
a
f
c
1 mm
1 mm 1 mm
h
e
Figure 2 | Cambrian arthropod eye SAM P43629, imaged using a Leica
MZ16FA relief-map stereomicroscope. a, Entire specimen showing the
positions of close-ups in c and f. b, Relief-map three-dimensional
reconstruction of a. c, Close-up of large ommatidial lenses in the bright zone,
with white line and numbers referring to the cross-section shown in e. d, Relief-
map three-dimensional reconstruction of c. e, Cross-section through four large
lenses indicated by the white line in c; numbers refer to individual lenses
represented by concavities. f, Close-up of small marginal lenses, with white line
and numbers referring to the cross-section shown in h. g, Relief-map three-
dimensional reconstruction of f. h, Cross-section through four small lenses
indicated by the white line in f; numbers refer to individual lenses represented
by concavities. Further pictures andimaging details are inSupplementary Fig. 2.
RESEARCH LETTER
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(ref. 18). The ommatidial gradient could have been achieved by two
different but non-exclusive developmental processes found in modern
arthropods. These are (1) a reduction in insulin/insulin-like growth
factor signalling fromthe centre tothe periphery, whichaffects cell size
19
;
and (2) the recruitment of a higher number of cells (mainly photorecep-
tors) ineachcentral ommatidium
20
. If these eyes grewinsize andomma-
tidial number during each instar by recruitment of new peripheral
ommatidial rows, as occurs in modern non-metamorphosing arthro-
pods
17
, then this too could have caused the older, central ommatidia to
consist of larger cells than the newly developed, marginal ommatidia.
The morphology of the visual surface and analogy to similar eyes in
living arthropods (Fig. 1d) indicate that each eye protruded laterally or
dorsolaterally, withthe bright zone directedanteriorlyor anterodorsally.
This bilaterally symmetrical arrangement generates binocular vision
(because the left and right bright zones have overlapping visual fields)
and also generates wide peripheral fields (because the smaller lenses
on the left and right eyes have largely complementary, rather than
overlapping, visual fields). However, confirmation of this inference
requires the discovery of articulated remains. Acute forward vision
and lower-resolution peripheral vision are typical of predators that
require excellent frontal vision for estimating distance and detecting
prey against complex backgrounds; they are also typical of fast-
moving organisms in which acute peripheral vision is precluded by
a high retinal angular velocity
21
.
The ommatidial lens facets in the Emu Bay Shale eyes are large
(60150 mm, Fig. 3b) but the lenses are not anomalously large given
the sizes of the eyes; relative sizes are broadly similar to those of
trilobites, living crustaceans and xiphosurans (Supplementary Fig. 4).
The association between the size and arrangement of ommatidia and
the level of ambient light has been quantified using the eye parameter
p (ref. 22). Owing to compression of the Emu Bay Shale eyes, p is
impossible to measure. However, p has been evaluated in detail in two
trilobites, Carolinites and Pricyclopyge
15
, with eyes broadly comparable
in both size and shape to the Emu Bay Shale eyes (that is, broadly
convex with hexagonally close-packed lenses). Carolinites genacinaca
has lenses up to 75 mm in size, similar to the ,6080 mm marginal
lenses of the Emu Bay Shale eyes, and Pricyclopyge binodosa has lenses
of up to 180 mm, comparable in size to the ,150 mm central lenses of
the EmuBay Shale eyes. Eyes of broadly equivalent shape andsize, with
similarly sized and packed lenses, could be expected to have similar
interommatidial angles and thus p values. If this extrapolation is valid,
the present eyes would have had p values comparable to the range
found across Carolinites and Pricyclopyge, that is, 2.138.31. Such
values are typical of taxa living in low-luminance environments
22
.
True compound eyes with lens-bearing ommatidia are restricted to
arthropods
23,24
. The complexity and large size of the Emu Bay Shale eyes
strongly indicate that they belongtoanactive arthropod, probably a large
predator. However, a definitive association with any particular taxon
must await articulated remains. In the Emu Bay Shale fauna, the only
sufficiently large arthropods knownare the trilobite Redlichia takooensis,
the stem-arthropod Anomalocaris and the bivalved arthropod Tuzoia
(ref. 11 andcitations therein). The eyes are clearly different inshape from
the seleniform, calcitic eyes of Redlichia and other trilobites. They also
seemtoo small to be referable to Anomalocaris. All of the eyes described
here are 79 mm in their longer diameter, suggesting that they came
from similarly sized adults. The expected eye diameter in adults of both
Anomalocaris species from the Emu Bay Shale is 23 times greater than
this, given the relative sizes of frontal appendages of anomalocaridids
from the Emu Bay, Burgess and Maotianshan Shales (with specimens
from the latter deposits having preserved eyes: ref. 25 and citations
therein). The large, unnamed Tuzoia species from the Emu Bay Shale
has stalked compound eyes that are ovoid to round and 69 mm in
diameter
11
: very similar to the fossil eyes described here. However, no
detailed structure of the visual surface is preserved in the articulated eyes
of Emu Bay Shale or Burgess Shale Tuzoia specimens
11,26
. Attribution of
the isolated eyes to Tuzoia would require a taphonomic explanation for
why disarticulated eyes are more commonly preserved and preserved in
finer detail. One possibility is that the fossils reported here are of pre-
viously shed corneas. The corneal surfaces of living arthropods detach
during ecdysis and remain loosely connected to the rest of the exuvia;
moulted corneas might be more prone to decay and thus more suscep-
tible to early diagenetic mineralization (in this case phosphatization)
than complete eyes attached to intact organisms.
The evolution of powerful vision is one of the most important cor-
relates
10
of the Cambrian explosion and has been proposed as a trigger
for this event
8
. However, although the overall shapes of eyes are known
for many Cambrian organisms
35
, intricate details of the visual surface
are known only for trilobites
27
and the tiny stem-crustacean cambro-
pachycopids, which have bizarre, proportionately huge and medially
fusedcompound eyes
9
. Inaddition, indistinct ommatidia are preserved
in a few Chengjiang fossils, including the non-biomineralized arthro-
pods Isoxys and Cindarella
28,29
. Isoxys inhabited both dim and bright
pelagic environments
28
whereas Cindarella probably inhabiteda bright
benthos
29
. The specimens described here represent the first microana-
tomical evidence confirming the view that highly developed vision in
a
Geological age (Myr ago)
b
Margin
Ordovician Cambrian
Ordovician Cambrian
100
200
300
400
0
0
1,000
2,000
3,000
4,000
5,000
6,000
N
u
m
b
e
r

o
f

o
m
m
a
t
i
d
i
a
O
m
m
a
t
i
d
i
a

s
i
z
e

(

m
)
Holochroal
Schizochroal
Abathochroal
Cambropachycopid
Emu Bay Shale eye
Trilobita
Holochroal
Schizochroal
Abathochroal
Cambropachycopid
Emu Bay Shale eye
Trilobita
Bright Zone
540 520 500 480 460 440
Geological age (Myr ago)
540 520 500 480 460 440
Figure 3 | Complexity of the Early Cambrian Emu Bay Shale eyes compared
to eyes in other early Palaeozoic taxa. a, b, Number of ommatidia (a) and lens
size (b) plotted against stratigraphic age for Cambro-Ordovician arthropods;
data in Supplementary Table 1. The Emu Bay Shale eyes have many more
ommatidia and much larger individual ommatidia than eyes in all other
Cambrian taxa. Trilobites are plotted according to eye type: schizochroal eyes
have relatively few, large lenses and are optically unusual compared to typical
compound eyes
27
.
LETTER RESEARCH
3 0 J U N E 2 0 1 1 | V O L 4 7 4 | N A T U R E | 6 3 3
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the Early Cambrian was not restricted to trilobites
8,10
. Furthermore, in
possessing more and larger lenses, plus a distinct bright zone, they are
substantially more complex than contemporaneous trilobite eyes,
which are often assumed to be among the most powerful visual organs
of their time
27,30
. Thenewfossils reveal that someof theearliest arthropods
had already acquired visual systems similar to those of living forms,
underscoring the speed and magnitude of the evolutionary innovation
that occurred during the Cambrian explosion.
Received 6 February 2011; accepted 1 April 2011.
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Supplementary Information is linked to the online version of the paper at
www.nature.com/nature.
Acknowledgements We thank P. and C. Buck for access to and assistance at the fossil
site; N. Schroeder, M. Gemmell, R. Atkinson, M. A. Binnie and numerous others
(Supplementary Table 3) for help with excavations and curatorial assistance;
A. Netting, P. Hudson and Adelaide Microscopy for imaging; D. Birch and G. Brock for
SEM-EDS analysis; A. Baonza and J. F. de Celis for discussions on arthropod eye
development; R. Fortey and A. Parker for comments and the Australian Research
Council (grant LP0774959), South Australian Museum, Spanish Ministry of Science
(RYC2007-00090 and grant CGL2009-07073), Beach Energy and Sealink Pty Ltd for
funding.
Author Contributions All authors contributed directly to excavation and interpretation
of fossil specimens, analysis and writing the paper. J.B.J., J.R.P. and M.S.Y.L. compiled
comparative eye data, M.S.Y.L. conducted the stereomicroscopy and J.R.P. conducted
the SEM-EDS analyses and digital photography.
Author Information Reprints and permissions information is available at
www.nature.com/reprints. The authors declare no competing financial interests.
Readers are welcome to comment on the online version of this article at
www.nature.com/nature. Correspondence and requests for materials should be
addressed to M.S.Y.L. (mike.lee@samuseum.sa.gov.au) or J.R.P.
(jpater20@une.edu.au).
RESEARCH LETTER
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Supplementary Information: Tables 1-3

SI Table 1. Number and average size of ommatidia for Cambrian and Ordovician arthropods (trilobites, cambropachycopids, and the Emu Bay
Shale eyes), as plotted in Fig. 3 (main text).

Species Clade Age
(Ma)
Eye
Type*
Number of
Ommatidia
Ommatidia
Lens
Diameter
(average, m)
Reference (1-30 cited in main text, 31-80 cited below)
Neocobboldia chinlinica Trilobita 520 A 84 20 Zhang & Clarkson 1990
16
, pls 3, 4, text-fig. 8; Gal et al. 2000
31
, fig. 1; Clarkson
et al., 2006
27
, fig. 4G, H
Shizhudiscus longquanensis Trilobita 520 A 95 37.5 Zhang & Clarkson 1990
16
, pls 1, 2
Pagetia prolata Trilobita 508 A 32 25 Jell 1975
32
, fig. 2E-H; Jell 1975
33
, pl. 12, figs 1-3
Pagetia ocellata Trilobita 507 A 20 20 Jell 1975
32
, fig. 4C, D
Pagetia sinesulcata Trilobita 507 A 59 33 Jell 1975
32
, fig. 2A-D; Jell 1975
33
, pl. 20, figs 1-3, 5
Pagetia thorntonensis Trilobita 506 A 46 35 Jell 1975
32
, fig. 3
Helepagetia argusi Trilobita 502 A 20 45 Jago 1972
34
, pl. 44, figs 4, 6, 9
Paradistazeris hubeiensis Trilobita 501 H 200 36 Peng et al. 2004
35
, pl. 21, fig. 11
Cedaria woosteri Trilobita 501 H 45 Hughes et al. 1997
36
, figs 4.2, 5
Catillicephalina
glasgowensis Trilobita 500 H 600 16 Bentley et al. 2009
37
, fig. 9H, I
Ctenopyge ceciliae Trilobita 491 H 150 21 Schoenemann et al. 2010
38
, text-figs 2, 3
Sphaerophthalmus alatus Trilobita 491 H 150 30 Clarkson 1973
39
, pl. 95, figs 1, 2
Peltura scarabaeoides Trilobita 491 H 180 40 Clarkson 1973
39
, text-fig. 4c
Ctenopyge (Eoctenopyge)
angusta Trilobita 491 H 200 50 Clarkson et al. 2003
40
, pl. 7, figs 4, 5
Sphaerophthalmus humilis Trilobita 491 H 200 50 Clarkson 1973
39
, pl. 94, fig. 6, pl. 95, figs 4-6
Jujuyaspis keideli Trilobita 488 H 1200 60 Aceolaza et al. 2001
41
, fig. 3
Ogygiocaris? cf. selwyni Trilobita 475 H 3000 62 Whittington 1966
42
, pl. 2, fig. 8
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Toletanaspis borni Trilobita 475 S 110 100 Henry et al. 1992
43
, fig. 2d
Oopsites hibernicus Trilobita 474 H 900 100 Fortey 1975
44
, pl. 34, figs 2, 5
Opipeuterella inconnivus Trilobita 474 H 2000 50 Fortey 1974
45
, pl. 14, figs 4, 6, 7
Carolinites genacinaca Trilobita 474 H 4000 75 Fortey 1975
44
, pl. 37, figs 7, 10, 12; McCormick & Fortey 1998
46

Telephina calandria Trilobita 473 H 900 80 Chatterton et al. 1999
47
, fig. 4.26
Psephosthenapsis glabrior Trilobita 471 H 5000 70 Fortey & Droser 1996
48
, fig. 11.8
Perischoclonus capitalis Trilobita 468 H 200 50 Whittington 1963
49
, pl. 22, fig. 7
Goniotelus kindlei Trilobita 468 H 220 40 Whittington 1963
49
, pl. 14, fig. 9; Whittington 1992
50
, pl. 4, fig. B
Kawina torulus Trilobita 468 H 350 60 Whittington 1963
49
, pl. 29, fig. 7
Bathyurellus nitidus Trilobita 468 H 850 50 Whittington 1963
49
, pl. 10, fig. 16
Apatolichas jukesi Trilobita 468 H 3500 30 Whittington 1963
49
, pl. 33, fig. 11; Whittington 1992
50
, pl. 4, fig. A
Carolinites rugosus Trilobita 466 H 1300 130 Fortey 1975
44
, pl. 36, figs 1, 7, 17
Retamaspis melendezi Trilobita 465 S 180 180 Hammann 1974
51
, pl. 2, fig. 17b
Pricyclopyge binodosa Trilobita 464 H 1200 180 Clarkson 1975
52
, pl. 2, figs 1-3, 5-8; McCormick & Fortey 1998
46
; Bruthansova
2003
53
, pl. 3, figs 9, 12

Asaphus cornutus Trilobita 464 H 3500 80 Clarkson in Whittington et al. 1997
54
, fig. 101.1; Fordyce & Cronin 1993
55
, fig. 6
Ormathops atava Trilobita 463 S 140 90 Henry et al. 1992
43
, fig. 1i
Telephina mobergi Trilobita 462 H 150 180 Ahlberg 1995
56
, pl. 3, figs 11, 12
Telephina bicuspis Trilobita 462 H 1100 180 Ahlberg 1995
56
, pl. 1, fig. 9; Bruton & Hyberget 2006
57
, fig. 1-O
Crozonaspis struvei Trilobita 462 S 184 270 Clarkson 1968
58
, pl. 1, figs 1-3
Microparia (Quadratapyge)
latilimbata Trilobita 461 H 350 70 Zhou & Zhou 2009
69
, fig. 2A, G
Sphaerexochus arenosus Trilobita 458 H 130 70 Chatterton & Ludvigsen 1976
60
, pl. 13, fig. 46
Remopleurides pattersoni Trilobita 458 H 5000 20 Chatterton & Ludvigsen 1976
60
, pl. 22, figs 12, 13
Calyptaulax callirachis Trilobita 458 S 130 220 Chatterton & Ludvigsen 1976
60
, pl. 16, fig. 34
Phacopidella hupei Trilobita 458 S 192 240 Nion & Henry 1966
61
, pl. 24, figs 1a, 1b, 1c, 2a, 2b, 2c.
Zeliszkella mytoensis Trilobita 458 S 200 210 Struve 1958
62
, pl. 1, fig. 7d
Kloucekia micheli Trilobita 458 S 280 210 Henry 1965
63
, pl. 1, fig. 9
Sphaerexochus pulcher Trilobita 456 H 75 60 Whittington & Evitt 1954
64
, pl. 21, fig. 2
Holia cimelia Trilobita 456 H 170 70 Whittington & Evitt 1954
64
, pl. 19, figs 3, 4; Whittington 1992
50
, pl. 4, figs C, D
Eomonorachus intermedius Trilobita 455 S 100 150 Ludvigsen & Chatterton 1982
65
, pl. 2, fig. 13
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Hispaniaspis (?) sp. indet. Trilobita 453 H 140 80 Fortey 1997
66
, pl. 10, fig. 4
Ovalocephalus ovatus Trilobita 453 H 160 80 Fortey 1997
66
, pl. 9, fig. 8
Telephina convexa Trilobita 453 H 1966 200 Fortey 1997
66
, text-fig. 3; Han 2001
67
, pls 1-4
Calyptaulax brongniartii Trilobita 453 S 225 280 Clarkson & Tripp 1982
68
, figs 4g, 5d; Clarkson et al. 2006
29
, fig. 4A, B
Achatella achates Trilobita 452 S 110 230 Ludvigsen & Chatterton 1982
65
, pl. 1, fig. 5
Sagavia chuanxiensis Trilobita 451 H 2000 60 Zhou & Zhou 2007
69
, fig. 2J
Flexicalymene cf. quadrata Trilobita 448 H 220 17 Clarkson 1975
52
, pl. 3, figs 9-11
Pseudogygites
latimarginatus Trilobita 447 H 40 Clarkson 1979
70
, pl. 1, fig. 6
Symphysops armata Trilobita 445 H 2500 140 Hammann & Leone 1997
71
, pl. 10, fig. 2
Mucronaspis mucronata Trilobita 445 S 120 210 Cocks & Fortey 1997
72
, pl. 1, fig. 9
Dalmanitina yichangensis Trilobita 445 S 230 360 Zhu & Wu 1983
73
, pl. 5, fig. 1
Goticaris longispinosa Cambropachycopidae 499 H 67 30 Haug et al. 2009
9
, pl. 5, fig. 1
Cambropachycope clarksoni Cambropachycopidae 499 H 149 40 Haug et al. 2009
9
, pl. 11, figs 3, 4
Emu Bay Eye margin Unknown 515 NA 3100 60 Current paper
Current paper Emu Bay Eye bright zone Unknown 150


* A = abathochroal, H = holochroal, S = schizochroal.
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Species Clade Eye radius
(r) (m)
Log10(r) Ommatidia
Diameter (d)
(m)
Log10(d) Reference (1-30 cited in main text, 31-80 cited below)
Polyphemus pediculus Crustacea 60 0.89 25 1.40 Nilsson & Oselius 1983
74
, Fig. 1
Leptodora kindtii Crustacea 210 1.16 30 1.48 Wolken & Gallik 1965
75
, Fig. 1
Gennadas sp. Crustacea 300 1.24 15 1.18 Meyer-Rochow & Walsh 1977
76
, p. 89, 91.
Thysanopoda tricuspidata Crustacea 550 1.37 28 1.45 Meyer-Rochow & Walsh 1978
77
, p. 61, Fig. 2b
Panulirus longipes Crustacea 5000 1.85 80 1.90 Meyer-Rochow 1975
78
, p. 440, 455.
Limulus polyphemus Xiphosura 6500 1.91 120 2.08 Fahrenback 1968
79
p. 279, fig. 1; Miller 1957
80
p. 426 gives similar ommatidia
diameter of 140 microns

Neocobboldia chinlinica Trilobita 120 1.04 22 1.34 Zhang & Clarkson 1990
16
, pl. 4, fig. 1a, b
Sphaerophthalmus alatus Trilobita 200 1.15 30 1.48 Clarkson 1973
39
, pl. 95, fig. 1
Perischoclonus capitalis Trilobita 410 1.31 50 1.70 Whittington 1963
49
, pl. 22, figs 2, 4, 6, 7
Apatolichas jukesi Trilobita 1060 1.51 30 1.48 Whittington 1992
50
, pl. 4, fig. A, pl. 96, fig. D
Oopsites hibernicus Trilobita 2100 1.66 100 2.00 Fortey 1975
44
, pl. 34, figs 2, 5
Calyptaulax brongniartii Trilobita 2530 1.70 280 2.45 Clarkson & Tripp 1982
68
, fig. 4d, g
Telephina convexa Trilobita 3370 1.76 194 2.29 Han 2001
67
, pl. 1
Pricyclopyge binodosa Trilobita 5350 1.86 135 2.13 Clarkson 1975
52
, pl. 2, figs 1, 2
Insects Insecta NA NA NA NA Wehner 1981
18
, p.309; see also Barlow 1952
81
.
Emu Bay Shale eye, margin 400 1.80 60 1.78 Current paper
Emu Bay Shale eye,
bright zone 400 1.80 150 2.18 Current paper

SI Table 2. Eye size and ommatidia size for a range of fossil and living aquatic arthropods. The data were combined with the data for terrestrial
arthropods in Wehner (1981 p. 309) and the combined data are plotted in SI Fig. 4.
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Organisation Personnel
Waterhouse Club (South Australian Museum) Carolyn and Trevor Ireland, Don and Margie Heylen, Mary-Lou and Antony Simpson,
Catherine Boros, Margaret Brown, Michael and Heather Clegg, Hank de Wit, Robert
Edwards, Sally Haynes, Robert Finlay-Jones, Suzanne Kent, Diana Laidlaw, Anne Levy,
Jan Lodge, Jan Perry, Bert and Barbara Prowse, Hamish Ramsey, Lyn Pederson, Rob
Searcy, John and Beth Shepherd, Ian Smith, Alun Thomas, Kathleen Cunningham, Diana
Watson, Ian and Mary Wilson
South Australian Museum Staff Natalie Schroeder, Dennis Rice, Mike Gemmell, Mary-Anne Binnie
University of Adelaide Staff Aaron Camens, Trevor Worthy, Jenny Worthy
Other Organisations / Private Paul and Carmen Buck, Ronda Atkinson, Glenn Brock, John Laurie, Pierre Kruse, Allison
Daley, Lars Holmer, John Ellice-Flint, Heather Catchpole, Richard Fortey, Roger Cooper,
Leonie Feutrill

SI Table 3. We thank the above associates and volunteers who have helped with excavations at Buck Quarry.
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References for Supplementary Information: Tables

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60. Chatterton, B.D.E. & Ludvigsen, R. Silicified Middle Ordovician trilobites from the South Nahanni River area, District of Mackenzie, Canada. Palaeontographica Abteilung A
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65. Ludvigsen, R. & Chatterton, B.D.E. Ordovician Pterygometopidae (Trilobita) of North America. Canadian Journal of Earth Sciences 19, 2179-2206 (1982).
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75. Wolken, J.J. & Gallik, G.J. The compound eye of a crustacean, Leptodora kindtii. Journal of Cell Biology 26, 968-973 (1965).
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Supplementary Information: Figures 1-4


SI Figure 1. Relief-map images of early Cambrian fossil eye (SAM P43629a)
showing three-dimensional preservation of fine surface structure. a, full visual
surface and b, close up of ommatidial lenses (area denoted by white box in a). Images
are rotated from top to bottom. Due to perspective there is no scale; see Main Text Fig.
1 and 2 for scales on planar images.
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SI Figure 2. Four eye specimens from the Emu Bay Shale, additional to the three
shown in Main Text Figure 1. All specimens positioned in the same orientation as
those in Figure 1. a, P45170a, b, P45913a and c, P44368b show most of the full ovoid
visual surface (at least in outline) but only patches of ommatidial lenses, d, P43445a is
a fragment of the visual surface showing well-preserved ommatidial lenses.
Counterparts are present for all specimens, but show little detail.
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SI Figure 3. SEM-EDS elemental maps of P43687. Each map shows the relative
abundance of each element; the brighter the colour, the more abundant the element.
Note that the visual surface of the eye contains high amounts of calcium (Ca) and
phosphorus (P), indicating a calcium phosphate composition. The matrix (shown top
left) contains elevated concentrations of oxygen (O), aluminium (Al) and silicon (Si),
representing the muscovite and other aluminosilicate clay minerals that constitute the
fossiliferous mudstones of the Emu Bay Shale.
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1.0 1.5 2.0
1.0
1.5
2.0
2.5
Bright Zone
Margin
Eye Size: Log10[(radius in m)]
O
m
m
a
t
i
d
i
a

s
i
z
e
:


L
o
g
1
0
(
d
i
a
m
e
t
e
r

i
n

!
m
)
Crustacea
Xiphosura
Insecta-Coleoptera
Insecta-Diptera
Insecta-Hymenoptera
Insecta-Lepidoptera
Insecta-Odonata
Trilobita-Holochroal
Emu Bay Shale eye
Trilobita-Abathochroal
Trilobita-Schizochroal
Emu Bay Shale eye


SI Figure 4. Ommatidial lens size compared to eye size, across fossil and living
arthropods. Data in Supp. Table 2; axis scales adopted from Wehner
18
(diameter of
lens vs square root of eye diameter, both on a log
10
scale in microns). Trilobites are
plotted separately according to eye type. The Emu Bay Shale eyes are broadly
comparable to Cambro-Ordovician trilobites and to living crustaceans and horseshoe
crabs.
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