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world's biggest trilobite--Isotelus Rex new species from the upper Ordovician of Northern Manitoba, Canada, The

Rudkin, David M

ABSTRACT-The largest known trilobite fossil. a virtually complete articulated dorsal shield of the asaphid Isotelus rex new species, has been
recovered from Upper Ordovician (Cincinnatian, Richmondian) nearshore carbonates of the Churchill River Group in northern Manitoba. At
over 700 mm in length, it is almost 70 percent longer than the largest previously documented complete trilobite. and provides the first
unequivocal evidence of maximum trilobite length in excess of one-half metre. Comparisons with other fossil and extant members of the phylum
suggest that in terms of maximum linear dimensions it was among the biggest arthropods ever to have lived. Sediments of the Churchill River
Group were deposited in an equatorial epeiric setting and the extremely large size of l. rex n. sp. thus marks a striking example of low-latitude
gigantism, in sharp contrast to the widespread phenomenon of "polar gigantism" in many modern marine benthic arthropods, Lack of extensive
epibiontic colonization of the exoskeletal surface and the presence of large distinctive trace fossils in the same unit suggest that l. rex n. sp. may
have been a semi-infaunal predator and scavenger that employed a shallow furrowing and probing mode of benthic feeding. The extinction of the
isotelines (and virtually the entire asaphide lineage) at the end of the Ordovician cannot be related to the near contemporaneous achievement of
exceptionally large adult size in some representatives. Failure to survive the terminal Ordovician extinction event was most likely a consequence
of a pelagic larval life-style that proved ill-adapted to the rapid onset of global climatic cooling and loss of tropical shelf habitats.

INTRODUCTION

THE TRILOBITES comprise a major class-level clade of extinct marine arthropods characterized by calcareous, multisclerite, dorsal exoskeletons.
The group has an excellent fossil record spanning some 300 million years-virtually the entire Paleozoic Era (Foote and Sepkoski, 1999). Despite
being constrained by a longitudinally trilobate body plan of tagmatized, serially arranged segments, trilobites attained a remarkable degree of
morphological diversity reflecting broad ecological adaptation within the marine biosphere (Wills et al., 1998; Fortey and Owens, 1999a,
1999b). Trilobites also varied greatly in size, with maximum reported holaspid (adult) lengths ranging from about 1 mm to about 700 mm
(Fortey and Whittington in Kaesler, 1997). The vast majority of trilobite species, however, were between 30 and 100 mm long at maturity
(Whittington in Kaesler, 1997), and only a small number of taxa are known to have exceeded 300 mm. Furthermore, the few previous literature
reports of trilobite lengths of 500 mm or more have been based not on complete articulated specimens, but on extrapolations from the dimensions
of partial exoskeletons or isolated sclerites. In this paper we report the discovery of a nearly complete trilobite fossil over 680 mm long that yields
a reconstructed exoskeletal length of more than 720 mm. This gigantic specimen, the holotype of Isotelus rex n. sp. (Fig. 1), provides the first
concrete evidence that trilobites could attain adult lengths of over one-half metre and represents the largest trilobite yet found.

STRATIGRAPHIC AND PALEOENVIRONMENTAL SETTING

The holotype and other specimens of Isotelus rex n. sp. documented herein were collected at a locality near the town of Churchill, Manitoba (Figs.
2, 3). The large exposure at this site is perhaps the most spectacular example of an exhumed Paleozoic rocky shore in the geologic record, and
happens to coincide with the modern coast of Hudson Bay (e.g., Norris and Sanford, 1969, fig. 10). Transgressive carbonates dipping gently
offshore were deposited over a boulder-strewn platform in front of an elongate ridge that protruded from the surrounding epeiric sea. The bedrock
and boulders are Precambrian Churchill Quartzite. Opinions on the age and stratigraphic affinity of the carbonates have changed over the years.
These deposits were thought to be Silurian by Nelson and Johnson (1966) and were assigned to the Severn River Formation by Sanford et al.
(1968). Norford (1971, addendum) reported Late Ordovician fossils and considered the strata to represent the Churchill River Group or Port
Nelson Formation. The beds were referred to as either Port Nelson or Severn River by Skinner and Johnson (1987), and as Port Nelson by Johnson
et al. (1988). in their important paleoenvironmental and paleoecological studies of the locality. Recently, Elias et al. (1999, Site 1) verified a
Late Ordovician (Cincinnatian, Richmondian) age for these strata and assigned them to the Churchill River Group on the basis of fossils and
lithology.

The stratigraphic interval in which I. rex n. sp. occurs (Fig. 3) consists of dolowackestone. The lower part (unit C) is comparatively rich in
bioclastic material such as pelmatozoan ossicles, and in quartz silt and sand; quartzite clasts up to 10 cm in diameter are present. Macrofossils
include colonial and solitary corals, nautiloid cephalopods, and aulacerid stromatoporoids. In the upper part (units D and E), Thalassinoides-type
burrow mottling is conspicuous, but macrofossils, including small fragmentary corals, are relatively rare. The upper bed of unit E, which yielded
the giant holotype of I. rex n. sp., contains rare evaporate-crystal molds. Microfossils in this bed include scolecodonts and conodonts. Among the
conodonts are the characteristic Richmondian species Oulodus rohneri and Pristognathus bighornensis; specimens suggestive of the subspecies
Pseudobelodina vulgaris ultima may be indicative of a late Richmondian age (Elias et al., 1999, fig. 2). The sediments of units C to E are
considered to have been deposited in a nearshore, shallow subtidal setting, with the energy level decreasing as water depth increased. The
conodont fauna represents a markedly shallow-water biofacies, but not the shallowest during this time interval. Paleogeographic reconstructions
place the area within 5 degrees of the Late Ordovician paleoequator (Scotese and McKerrow, 1991).

Thus far, complete or nearly complete specimens of I. rex n. sp. have all been found dorsal side up. The holotype specimen was oriented
approximately perpendicular to, and facing away from, the paleoshoreline, probably very close to where it lived, died, and subsequently became
interred. Isotelus rex n. sp. occurs near the very top of the Late Ordovician sequence in the Churchill area, where overlying strata of the lower
Severn River Formation are dated as Early Silurian (Llandovery, late Rhuddanian) (Elias et al., 1999). Isotelines (indeed, the entire family

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Asaphidae) did not survive the terminal Ordovician extinction crisis and the Churchill River Group fossils represent some of the last members of
a widespread and moderately speciose phyletic lineage whose cryptic origins extend back into the Cambrian Period (Fortey and Owens in Kaesler,
1997).

SIZES OF LARGE TRILOBITES

The giant trilobite documented here represents a new species of the widely distributed Ordovician asaphid genus Isotelus. Large representatives of
Isotelus occur elsewhere in the Late Ordovician succession of North America and some of the earliest described species were considered to be
among the biggest trilobites then known. Indeed, several specific epithets, including those of I. gigas Dekay, 1824 (type species; Fig. 4.1) and I.
maximus Locke, 1838, were coined in reference to their comparatively large size. Hansen (1985, 1989) has reported rare articulated specimens
of I. brachycephalus Foerste, 1919 from Ohio with lengths between 350 and 410 mm (see also Fig. 4.2).

Recently authenticated examples of complete, giant non-asaphide trilobites include an olenelline, Cambropallas (230 mm) and a redlichiine,
Acadoparadoxides (390 mm; Fig. 4.3), both from the Cambrian of Morocco (Geyer, 1993). There have been persistent reports in the literature of
lichid trilobites with lengths exceeding 500 mm, but these were based on fragmentary remains or large disarticulated sclerites preserved in
shaly lithologies susceptible to compaction (Reimann, 1942; Rabano, 1989; Whittington in Kaesler, 1997). For example, a widely cited
maximum holaspid length of 660 mm for Uralichas hispanicus (Ordovician; Iberian Peninsula) is apparently based on summed measurements of
individual large cephalic, thoracic, and pygidial elements (Rabano, 1989); moreover, at least 20 percent of the total sagittal length can be
accounted for by the narrow, blade-shaped median spine extending from the posterior pygidial margin (Fig. 4.4). To compound the difficulty of
obtaining accurate measurements, illustrated sclerites of Uralichas often show considerable tectonic deformation (Rabano, 1989, figs. 3, 6),
rendering some dimensions suspect. Similarly, restored lengths of up to 600 mm reported for the elaborately spinose lichid Terataspis (Devonian;
eastern North America) were also determined from dimensions of disarticulated sclerites which are frequently compacted and distorted
(Reimann, 1942). Whittington (in Kaesler, 1997) has summarized several other records of large trilobites.

Prior to this report, the largest confirmed complete trilobite specimen was a fully articulated isoteline measuring 430 mm in length (Johnson et
al., 1988, fig. 13a; Whittington in Kaesler, 1997; Figs. 4.5, 6.5). It is from the same locality as the much bigger holotype specimen of Isotelus rex
n. sp., and is herein assigned to the same species.

Isotelus is a moderately effaced (smooth) asaphid typically lacking terminal spines or prolongations. and dimensions of the exoskeleton therefore
closely approximate actual body size. The giant holotype specimen of I. rex n. sp. (Figs. 1, 6.1-6.4) was found in a competent carbonate unit and
shows little evidence of distortion or compaction. About two-thirds of the exoskeleton is retained as a thin (approximately 0.5-2 mm thick)
calcareous layer adhering to the convex internal mould (Fig. 6.3). All dorsal sclerites are in close articulation and there are no gaps along
cephalic sutures (Fig. 1). The fossil is clearly an undisturbed carcass of a dead animal and not an exuvium. The dimensions of the fossil are
probably not significantly different from those of the living animal (exclusive of unpreserved appendages) and at over 700 mm long, 400 mm in
maximum width (across the cephalon), and approximately 70 mm in height (at the posterior midpoint of the cephalon), it is the largest known
trilobite (Fig. 4.6).

ACKNOWLEDGMENTS

Young and Elias acknowledge individual grants in support of this project from the Natural Sciences and Engineering Research Council of Canada.
Young also received funding from the University of Manitoba and the Manitoba Museum Foundation. Rudkin is grateful for continuing grant
support from the Royal Ontario Museum Foundation. Valuable field assistance was rendered by J. D. Klapecki and D. L. Wright (MMMN), and N.
J. Aime, S. H. Margerison and C. J. Moffat (University of Manitoba). We thank R. Barrow (MMMN) for photographing the holotype specimen and
K. David and P Fenton (ROM) for production of the outline drawings in Figures 4 and 5. The photograph of GSC 85292 was generously provided
by the late T. E. Bolton. B. D. E. Chatterton and an anonymous referee are thanked for careful and constructive reviews of the manuscript.

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DAVID M. RUDKIN,1 GRAHAM A. YOUNG,2 ROBERT J. ELIAS,3 AND EDWARD P DOBRZANSKI2

1Department of Palaeobiology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario M5S 2C6, Canada, ,

2The Manitoba Museum, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2, Canada, and

3Department of Geological Sciences, The University of Manitoba, Winnipeg, Manitoba R3T 2N2, Canada

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