A New ?

Lamellipedian Arthropod from the Early Cambrian Sirius Passet Fauna

of North Greenland
Author(s): Linda Lagebro , Martin Stein , and John S. Peel
Source: Journal of Paleontology, 83(5):820-825. 2009.
Published By: The Paleontological Society
DOI: http://dx.doi.org/10.1666/09-011.1
URL: http://www.bioone.org/doi/full/10.1666/09-011.1

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J. Paleont., 83(5), 2009, pp. 820–825
Copyright ’ 2009, The Paleontological Society
0022-3360/09/0083-0820$03.00

A NEW ?LAMELLIPEDIAN ARTHROPOD FROM THE EARLY

CAMBRIAN SIRIUS PASSET FAUNA OF NORTH GREENLAND
LINDA LAGEBRO,1 MARTIN STEIN,1,2 AND JOHN S. PEEL1
1
Department of Earth Sciences, Palaeobiology, Uppsala University, Villavägen 16,
SE-752 36 Uppsala, Sweden, ,linda.lagebro@spray.se., ,john.peel@pal.uu.se.; and 2present address: Museum of Evolution,
Uppsala University, Norbyvägen 16, SE-752 36 Uppsala, Sweden, ,martin.stein@evolmuseum.uu.se.

INTRODUCTION SIRIOCARIS TROLLAE new genus and new species

arthropod described herein is Figures 1–5
T HE NON-MINERALIZED
derived from the Sirius Passet fossil conservation deposit
of North Greenland (82u47.69N, 42u13.79W), the oldest
Diagnosis.—Arthropod of about 100 mm length, sub-
elliptical habitus. Parabolic head shield, weakly trilobate,
locality with exceptional preservation of soft tissues known one fifth of total length. Trunk of about twenty segments; axis
from the Cambrian of Laurentia (Cambrian Series 2, Stage 3; wide, tergal pleurae divided into inner and outer portions.
Nevadella Zone). As such, it is broadly contemporaneous Inner portion with articulating flange anteriorly. Outer
with the Chengjiang fauna of China (Hou et al., 2004) and portion with prominent pleural spine; falcate in anterior seven
some 10 million years older than the Burgess Shale fauna of to eight tergites, sickle shaped in posterior segments. Tail
British Columbia. The Sirius Passet fauna was first docu- shield small, consisting of axial portion and single plate.
mented by Conway Morris et al. (1987) and its geological Antennae long and stout, multiarticulate. Three pairs of
setting is discussed by Babcock and Peel (2007). In addition postantennal limbs in head. All postantennal limbs biramous,
to the nevadiid trilobite Buenellus higginsi Blaker, 1988, the with medially biserially spinose basipod, robust endopod, and
fauna is dominated by non-mineralized arthropods (Budd, exopod with large, blade-shaped distal portion.
1993, 1995, 1997, 1999; Williams et al., 1996; Taylor, 2002). Description.—(a) Dorsal morphology: General habitus sub-
Other finds include sponges (Rigby, 1986), a lobopod (Budd elliptical, about 2.5 times as long as wide; maximum width at
and Peel, 1998), the earliest annelids (Conway Morris and third and fourth tergites (Fig. 2). Total length varies between
Peel, 2008) and articulated halkieriids (Conway Morris and 100 and 110 mm in the most nearly complete specimens. Head
Peel, 1990, 1995), but most of the assemblage awaits
description.
Siriocaris trollae gen. et sp. nov. has long, multiarticulate
antennae, like lamellipedian arthropods, and the dorsal
morphology shows particular similarities with trilobites and
helmetiids. Some 14 specimens have been studied. Due to
their relatively large size, most are broken, but there is no
evidence of disarticulation. Six specimens preserve only
tergites, head shield, and tail, mostly embedded in dorsoven-
tral orientation, with the tergites lying flat on the bedding
plane. Five specimens preserving limbs are embedded in
oblique orientation, with the limbs of one series extending
from under the tergites and those of the other, except the
most anterior, either folded under the tergites or not
preserved.
In the description below, terminology follows Stein et al.
(2008) for ventral morphology, except that the term antenna is
applied for the anteriormost appendage, the antennula in
crustacean terminology. Terminology for dorsal morphology
follows Whittington (1997). Type specimens are deposited in
the Geological Museum (State Museum of Natural History),
Copenhagen, Denmark (MGUH prefix). GGU indicates
collections of the Geological Survey of Greenland (now
Geological Survey of Denmark and Greenland), Copenhagen,
Denmark.

SYSTEMATIC PALEONTOLOGY
ARTHROPODA von Siebold and Stannius, 1854
?LAMELLIPEDIA Hou & Bergström, 1997
SIRIOCARIS new genus
Type species.—Siriocaris trollae, sp. nov. (by monotypy).
Diagnosis.—As for species.
Etymology.—Sirio from Sirius Passet, and caris, Latin for FIGURE 1—Reconstruction of Siriocaris trollae gen. et sp. nov. Right
‘‘shrimp’’. half of eighth tergite omitted to show articulating flange.

820
 LAGEBRO ET AL.—AN EARLY CAMBRIAN ARTHROPOD FROM GREENLAND 821

FIGURE 2—Siriocaris trollae gen. et sp. nov. 1, Holotype MGUH 28970, arrows point to folds marking the raised axis of the head; light from upper
left; 2, details of tergites and tail, anteriormost tergites and head shield not preserved; MGUH 28973; light from upper left. Abbreviations: af, axial
furrow; ant, antenna; ba, basipods; cs, head shield; en, endopod; ex, exopod; fl, flange; ip, inner portion of pleurae; op, outer portion of pleurae; psp,
pleural spines; tg, tergite tl, tail; tla, axial portion of tail; tlp, tail plate.

shield is approximately 1/5 of total body length, parabolic in
outline. Posterolateral corners form acute angle. Weakly
trilobate.
Trunk consists of about 20 segments. First tergite roughly
same width (tr.) as head shield. Width increases to third
tergite, both width (tr.) and length (sag.) decrease posterior to
fourth tergite. Posteriormost tergite about 1/4 to 1/5 the
length (sag.) of first tergite. Axis occupies about half the
tergal width throughout trunk. Pleurae divided into inner and
more oblique outer portion that forms pleural spine
(Fig. 2.2). Anterior margin of tergites is straight throughout
inner portion and forms distally a flange stacked under
posterior border of preceding tergite (Fig. 2.2); distally,
anterior margin curves back into spine. Pleural spines are
falcate in anterior seven to eight tergites, grading into sickle
shaped in more posterior tergites. Length of pleural spines
increases gradually towards posterior; spines of first tergite
extend about 1/4 the length of the second tergite; spines of
thirteenth to sixteenth tergites each extend back through the
length of two following tergites. Pleural spines of seventeenth
to posteriormost tergites decrease in size. Width (tr.) of outer
pleural portion increases relative to that of the inner portion
across trunk toward posterior; posteriormost tergite has only
outer portion. Posterior margin straight axially, curves back
into pleural spines abaxially. Axially, each tergite overlaps
the following tergite posteriorly by 1/6 to 1/5 the tergal length
(sag.).
Tail piece small; 1/8 to 1/7 the width of maximum width of
trunk. Laterally flanked by pleural spines of five posterior-
most tergites. Consists of plate extending posterior to a short,
raised axial portion that terminates in parabolic shape
(Fig. 2.2). Lateral borders of plate straight anteriorly,
extending backward and outward; no pleural portions lateral
to raised axial portion.
(b) Ventral morphology: Head carries uniramous antennae
and three pairs of biramous limbs. Eyes, hypostome, and
position of mouth and limb insertions unknown. Sturdy but
flexible antennae protrude from under head shield at
anterolateral margin, separated by 1/3 the head shield width
(Fig. 2); their minimum length at least 2/3 the total length of
body; composed of short cylindrical articles carrying setae
medially (Fig. 3.2); number of articles unknown. First
postantennal limb protrudes from anterolateral margin of
head shield close to antennae (Fig. 4.1). All postantennal
limbs biramous, more or less undifferentiated except in size;
size increases from first limb to anterior thoracic limbs
(Fig. 4.1). Length of limbs about 4/5 of tergite width (tr.).
Basipod about 1/4 to 1/3 of limb length, biserially spinose
medially (Fig. 4.1). Endopods robust, number of podomeres
unknown. Proximal podomeres massive, distal ones narrow
rapidly; distal podomere hook-like, possibly flanked by spines
(Fig. 4.3). Setose endites unknown. Exopods of almost equal
length as endopods, situated adjacent to lateral edge of
endopods (Fig. 4.1, 4.3); articulation with basipod and
endopod unknown. Outline of distal part of exopod leaf-
shaped, more rounded in anterior exopods; setation unknown.
Etymology.—For Trolla.
Holotype.—MGUH 28970.
Other material.—A total of 13 specimens from the same
locality, GGU collection 340103; MGUH 28971–28983.
822 JOURNAL OF PALEONTOLOGY, V. 83, NO. 5, 2009
FIGURE 3—Siriocaris trollae gen. et sp. nov. 1, MGUH 28977, showing the maximum preserved length of the antenna; light from upper left; 2,
MGUH 28970 (holotype), detail of proximal part of antenna, arrows point to article joints; light from left. Abbreviations other than in previous figures:
ed?, possible enditic protrusions; s, setae.
Occurrence.—Peary Land, central North Greenland; base of
the Buen Formation (Cambrian Series 2, Stage 3, Nevadella
Zone).
Discussion.—Tergal boundaries are often obscure in the
anteriormost and particularly in the posteriormost segments,
which makes exact determination of their number difficult. The
status of the head shield with regards to trilobation is unclear.
The trunk is trilobate, with a raised axis and lowered pleurae. In
some specimens, a hint of an axial furrow is present posteriorly
(Fig. 2.2), but it is shallow and ill-defined. On the head shield,
there are distinct folds concentric to the shield margin, setting
off what seems to be an axial region (Fig. 2, arrows), most
apparent anteriorly. However, in the absence of distinct axial
furrows, and considering the occurrence of compactional folds
from flattening, the assumption of trilobation is tentative. The
division of the pleurae into inner and outer portions is indicated
by a slight difference in relief (Fig. 2.2).
Only the anterior portions of the tail piece are preserved,
showing the raised axial region and the anterior part of the
plate with its lateral margins extending backward and
somewhat outward from behind the axial piece (Fig. 2.2).
The outline of the plate is unknown. It is not known if the tail
carried limbs. Preservation is too poor to determine if the axial
portion consisted of one or more axial rings. The position of
the anus is unknown.
No ventral features of the head, such as limb insertions, the
mouth, or the hypostome are preserved. It is possible that the
ventral side of the head also contained visual organs, but there
is no preserved evidence.
The number of antennal articles can not be determined; the
distal portions are not preserved, and even in the proximal
portions, fidelity of preservation is only sufficient to detect
some article joints (Fig. 3.2, arrows). Setae on the articles are
rarely preserved (Fig. 3.2) and their position in relation to
article joints is not known. Only the proximal portions of setae
are preserved, so that their length cannot be determined.
Limb insertions cannot be traced, but impressions of the
basipods give an indication of their position. MGUH 28970
shows impressions of three basipods under the head shield,
and corresponding limbs can be seen distal of it (Fig. 4.1).
Basipods are known only from impressions under the
tergites and head shield, usually as elongate concave areas; in
one case the spines of the median edge are impressed
(Fig. 4.1). The two rows of spines seem to be set wide apart
proximally, about 1/3 to 1/2 the tergite length, and converge
distally. Of the endopods, mostly the more distal portions are
preserved. The proximal portions are usually covered by the
tergites and the distal portions are often partially covered by
the exopods. Few details of the endopods can be established,
but pivot joints are preserved in MGUH 28982 (Fig. 4.2). It is
not known if the articles had any kind of setation or medial
enditic projections. MGUH 28977 shows imprints of medio-
distal projections that may be spines or enditic projections
(Fig. 3.1), but these impressions may also be traces of a frayed
exopod margin overlying the endopod. Only the distal
portions of the exopods protruding from under the tergites
are accessible, consisting of a blade-like structure without any
setae preserved. Proximally, close to the tergites, the outer end
of the exopod curves inward, indicating that the proximal part
may be of different morphology (Fig. 4.4). The preserved
distal portions may correspond to the ‘‘distal lobe’’ (Ramsköld
and Edgecombe, 1996) of trilobite-like arthropods which in
 LAGEBRO ET AL.—AN EARLY CAMBRIAN ARTHROPOD FROM GREENLAND 823

FIGURE 4—Siriocaris trollae gen. et sp. nov. 1, MGUH 28970 (holotype), details of limbs showing basipod spines (ba1–8), arrows point to podomere
boundaries; light from lower left; 2, MGUH 28982, detail of endopod showing pivot joints (arrows); light from upper light; 3, MGUH 28981, detail of
distal parts of endopods; light from upper right; 4, MGUH 28977 detail of exopods; light from lower left.
824 JOURNAL OF PALEONTOLOGY, V. 83, NO. 5, 2009
FIGURE 5—Siriocaris trollae gen. et sp. nov, putative soft tissue
preservation. 1, MGUH 28977, arrow points to putative muscle tissue;
light from lower left. 2, MGUH 28981; light from upper left. 3, MGUH
28982; light from lower left.
some species, e.g., Naraoia spinosa Zhang and Hou, 1985
(Zhang et al., 2007), may be the dominant part of the exopods.
In Kuamaia lata Hou, 1987 the distal lobe has a somewhat
comparable shape; the supposed subdivision (Hou and
Bergström, 1997) is considered an artifact (Edgecombe and
Ramsköld, 1999). Setae are preserved neither on the blade-like
accessible part nor proximal to that.
Biserially arranged clover-shaped pits occur in the axial
region of MGUH 28970 and MGUH 28983 (Fig. 2.1); in one
specimen there are slit-like pits perpendicular to the axis
(Fig. 5.1). The clover-shaped pits often contain striations or
stacked sheets (cf. ‘‘planar elements’’ of Butterfield, 2002) that
are subradially arranged or parallel to the axis. Similar
structures occur frequently in exceptionally preserved arthro-
pod fossils and are commonly interpreted as midgut diverti-
culae, preserved through early permineralization (Butterfield,
2002). The only possible trace of the alimentary canal occurs
in specimen MGUH 28981. It is a more or less continuous
sagittal hollow with raised edges under the fifth to eighth
tergites (Fig. 5.2). MGUH 28982 shows a series of exsagittal
brush-like structures. These are bundles of crescentic grooves,
fanning out distally, arching outward and backward (Fig. 5.3).
They are of high relief, carving deep into the tergites, and
surrounded by a distinct yellow-orange stain. Their origin is
clearly segmental, with one bundle per tergite, but the distal
parts traverse tergal boundaries. The type of preservation
suggests an internal structure. The exsagittal position renders
interpretation as gut diverticulae unlikely, but could better
match extrinsic limb musculature. Muscle preservation in
Pambdelurion whittingtoni Budd, 1997 has been described from
the Sirius Passet fauna (Budd, 1998). Similarities between the
structures, however, are tenuous. Structures more resembling
the putative muscle tissue in P. whittingtoni are found in the
central part of the axis of MGUH 28977 (Fig. 5.1, arrow) or
the striation or stacked sheets in the putative gut diverticulae.
Among arthropods from the Sirius Passet fauna, Siriocaris
trollae is readily distinguished from Buenaspis forteyi Budd,
1999 by the relatively longer form with a tapering trunk of
about 20 segments and absence of a large pygidium.
Kleptothule rasmusseni Budd, 1995, differs in a triangular
head with ‘anterior spikes’, dorsal structures interpreted as
eyes, a considerably narrower shape, dorsal furrows on the
tergites, and large, caudal area of possibly ankylosed tergites
(Budd, 1995).
Flagelliform antennae such as those in Siriocaris trollae are
found in Lamellipedia (Trilobita and closely related ‘trilobi-
tomorph’ taxa). Exopod morphology may set S. trollae aside
from trilobites or other lamellipedians which commonly have
a bipartite exopod with a proximal portion carrying lamellar
setae and a distal, lobe-like portion with a setal fringe (e.g.,
Edgecombe and Ramsköld, 1999). It is not clear if such an
exopod morphology actually was present in S. trollae; the
proximally inward curving outer edge of the exopod (Fig. 4.4)
may suggest that the visible part indeed is only the distal lobe-
like part. Further comparisons face the dilemma that a setal
fringe is at least not preserved, and there is no compelling
evidence of lamellar setae. Some aspects of the postcephalic
dorsal morphology, such as the edge-edge articulation with
wider tergal overlap in the axis and a more or less horizontal,
hinged, inner portion of the pleura, are in accord with
trilobites and helmetiids (Edgecombe and Ramsköld, 1999).
Falcate to sickle-shaped pleurae with articulating flanges and
a small, plate-like tail clearly set S. trollae apart from
helmetiids and other non-trilobite lamellipedians but are
reminiscent of olenelline trilobites. However, the absence of
dorsal eyes, a clearly defined glabella, and dorsal furrows
other than the vague axial furrow and further articulating
devices, as well as the non-mineralized cuticle, clearly
differentiate S. trollae from olenellines or other trilobites.
Further comparisons with lamellipedians will require more
information on the limb morphology of S. trollae.
ACKNOWLEDGMENTS
Expeditions to Sirius Passet were made possible by financial
support from the Carlsberg Foundation (Copenhagen) and
National Geographic Society (Washington D.C.) to J. S. Peel
and S. Conway Morris. The logistic framework was partly
coordinated by the Geological Survey of Greenland and the
Danish Polarcenter. Financial support from the Swedish
Research Council (Vetenskapsrådet) to J. S. Peel is gratefully
acknowledged. We thank G. D. Edgecombe and B. S.
Lieberman for constructive reviews of the manuscript.
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ACCEPTED 13 MAY 2009