An Ibexian (Early Ordovician) trilobite faunule from the type section of the

Rabbitkettle Formation (southern Mackenzie Mountains, Northwest Territories)

BRIANR. PRATT
Department of Geology, University of Toronto, Toronto, Ont., Canada M5S 1Al
Received October 23, 1987
Revision accepted February 18, 1988

The Rabbitkettle Formation is a widespread lower Paleozoic unit in the northern Cordillera. A trilobite faunule collected
from the middle part of the type section and described here documents an early Ibexian (Early Ordovician) age for this interval
Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by University of Saskatchewan on 05/07/14

and provides the first biostratigraphic reference point for the type section. This faunule is dominated by the agnostoid Micra-
gnostus, the leiostegiid Leiostegium, and the remopleuridid Kainella. These genera and rarer taxa have North American, South
American, and Austral-Asian affinities. A new species, Kainella eleutherolf, is established.

La formation de Rabbitkettle est une unit6 du PalCozoique infkrieur largement dpandue dans la Cordilkre du nord. Une
faunule de trilobites, CchantillonnCe dans la partie moyenne de la section type et dtcrite ice, donne un Pge Ibexien prCcoce
(Ordovicien prCcoce) pour cet intervalle et foumi le premier point de kfCrence biostratigraphique pour la section type. Cette
faunule est dominee par l'agnostolde Micragnostus, le leiostbgiide Leiostegium et le kmopleuridide Kainella. Ces genres et
taxa plus rares ont des affinitks nord-americaines, sud-americaines et austral-asiennes. Une nouvelle espkce, Kainella
eleutherolf, est Ctablie.

Can. J. Eanh Sci. 25, 1595 - 1607 (1988)

Introduction in this area. By current convention (North American Commis-

The Rabbitkettle Formation was formally proposed by sion on Stratigraphic Nomenclature 1983), these shortcomings
Gabrielse et al. (1973, p. 48) as a name for a thick, distinctive make this locality a poor choice for a unit stratotype.
For personal use only.

sequence of interbedded silty limestones and limy siltstones
exposed in the southern Mackenzie Mountains of northwestern
Canada. It replaced unit 9 of Blusson (1968) and has a desig-
nated type area near the headwaters of Rabbitkettle River and
Flat River (Fig. 1). Although no fossils were found in the type
area, Blusson (1968) presumed the unit to be Middle to
possibly Late Cambrian in age, whereas Gabrielse et al. (1973)
suggested a Late Cambrian and Early Ordovician age because
it overlies with angular unconforrnity the Lower Cambrian
Sekwi Formation and is in turn unconformably overlain by
upper Middle Ordovician graptolitic shale near Flat River.
Usage of the name Rabbitkettle Formation has extended
northwestward along the entire Mackenzie Mountains as far as
Arctic Red River (Tipnis et al. 1978; Gordey 1979, 1980;
Gordey et al. 1981; Landing et al. 1980; Fritz 1981; Ludvigsen
1982a, 1982b; Cecile 1982), and the formation has been corre-
lated with the Kechika Group of the northern Rocky Mountains
and Cassiar Mountains of northern British Columbia and adja-
cent Yukon Territory (Gabrielse et al. 1973, p. 51; Fritz
1985). The Rabbitkettle thus represents a lithologic unit that is
widely distributed along the northern Cordillera as a facies
transitional from shallow-water carbonates to the east and
basinal shales to the west.
Precise correlations, however, have been hampered because
the age of the Rabbitkettle is only broadly constrained by stra-
tigraphic relationships, and firm faunal control has been
absent. This paper describes the first and thus far only fossils
recovered from the Rabbitkettle Formation at its type area.
Stratigraphic section
The type area of the Rabbitkettle Formation is a discontinu-
ous series of measured sections (sections 14 and 15 of Blusson
1968) along a mountainside southwest of the Flat River,
18-20 krn southeast of Tungsten, Northwest Temtories. The
unit is over 1500 m thick, but its base and top are not exposed
Pnnted in Canada I Imprim.6 au Canada
The Rabbitkettle at its type area consists mainly of thin-
bedded, laminated argillaceous and calcareous siltstone and
lenticular- and nodular-bedded silty lime mudstone, with scat-
tered medium to thick beds of burrow-mottled silty lime mud-
stone (Fig. 2). In plan view, many of the lime mudstone
nodules are interconnected, resembling horizontal 77zalassi-
noides burrow systems; small, spar-filled burrows are locally
preserved in the nodules. Short vertical spreite burrows and
squashed trilobite fragments are occassionally seen in the lami-
nated siltstone. Only one thin, lenticular bed of silty bioclastic
grainstone was observed; this bed contained the trilobites
described here and pelmatozoan ossicles (Fig. 3). Trilobite
remains are occasionally present in the lime mudstone beds,
but their primary fibrous calcite skeletons have recrystallized
to equant micrite and microspar, too similar to the matrix to
permit recovery by splitting.
Tectonic deformation from west-east compression has
affected the entire section, and most of the trilobites are dis-
torted. Lenticular and nodular lime mudstone beds have been
formed into lumpy nodules up to about twice their original
thickness. The 1500 m of Rabbitkettle measured by Blusson
(1968) is therefore an exaggeration of true thickness, perhaps
by as much as one third.
The Rabbitkettle lithologies at the type area suggest a rela-
tively deep depositional environment; the near absence of
grainstone horizons suggests that it was mostly below storm
wave base, i.e., below 100 m depth for a location facing the
open ocean. The absence of slumped beds and obvious trans-
ported units such as turbidites and debris-flow conglomerates
indicates that the depositional slope was low and that the over-
all environment may be characterized as a deep shelf. The
trilobite fauna recovered from the single grainstone bed is
interpreted as indigenous (that is, a rare, in situ product of win-
nowing and concentration) because of the lack of associated
coarse-grained allochthonous beds, the lack of abrasion of the
fossils, and the presence of scattered trilobite fragments in
 1596 CAN. J. EARTH SCI. VOL. 25, 1988
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For personal use only.
FIG. 1. (A) Map of northwestern Canada showing location of the Flat River area (B). (C) Topographic map of the type area of the Rabbitkettle
Formation shows the location of Blusson's (1968) sections 14 and 15 and the trilobite locality.
interbedded siltstones and lime mudstones.
Blusson (1968, p. 14) reported finding unidentified trilobite
fragments from the middle part of the formation at a locality 12
krn to the west of the type section. The fossils described here
also came from the middle part, near the top of Blusson's
(1968) section 14 at 61°47'N, 127O57'W.
Age and composition of the faunule
The trilobite faunule is dominated by three genera, Micra-
gnostus, Leiostegium, and Kainella (Fig. 4; Appendix); in
addition, rare components include Angelina, Apatokephalus,
Perischodory, and Gymnagnostus?. Although a trilobite bio-
facies scheme has not yet been formulated for rocks of this
age, lithostratigraphic observations suggest that this faunule
represents a deep-shelf association of taxa tolerant of some-
what cooler water than that of the adjacent carbonate platform
of the Broken Skull Formation. Most of these genera are world
wide in their distribution, but the Rabbitkettle faunule specifi-
cally links those of shallow-water carbonate strata of the Great
Basin with those in colder water, noncarbonate strata of Argen-
tina, northwest Europe, Tasmania, and central Mexico.
Leiostegium manitouensis is the most characteristic trilobite
of Zone D of the Utah-Nevada Ibexian trilobite zonation
(Ross 1949; Hintze 1952; Ross et al. 1982). The abundance of
this species in the Rabbitkettle collection indicates a Zone D
age for this faunule. This is supported by the presence of
Apatokephalus finalis, which also occurs in Zone D strata in
Nevada (Hintze 1952). This zone, termed the Leiostegium -
Kainella Zone by Hintze (1952), also yields a species of
Kainella in Nevada. This genus is represented in the Rabbit-
kettle by Kainella eleutherolji. In northwestern Argentina
there are two species: Kainella conica, from the Parabolina
argentina Zone of the lower part of the lower Tremadocian,
and Kainella meridionalis, which characterizes the Kainella
meridionalis Zone of the upper part of the lower Tremadocian
(Hamngton and Leanza 1957). Angelina hyeronimi occurs in
the Kainella meridionalis Zone of Argentina as well as in cen-
tral Mexico. The presence of this species in the Rabbitkettle
collection suggests that Zone D correlates with the Kainella
meridionalis Zone of the lower Tremadocian as defined in
Argentina. Leiostegium douglasi, a species very close to Leio-
stegium manitouensis, also occurs in this zone. No species
from the Rabbitkettle collection are common to the Tremado-
cian of Great Britain. However, Apatokephalus serratus,
which occurs in the early upper Tremadocian Shumardia
pusilla Biozone of England and Wales (Rushton in Whittington
et al. 1984; Thomas et al. 1984), also occurs in the Kainella
meridionalis Zone and in the upper Tremadocian of Argentina.
This suggests that the Rabbitkettle collection is approximately
correlative with the middle Tremadocian of Great Britain.
The presence of the collection in the middle part of the
Rabbitkettle suggests that the formation in its type area ranges
upward into the later Ibexian at least and downward well into
the Late Cambrian. It is not known, however, if the lower part
is correlatable with the Rockslide Formation defined farther
north.
Systematic paleontology
Repository
Illustrated and type specimens are housed at the Royal
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For personal use only.
FIG.2. Lithology of the Rabbitkettle Formation. (A) Outcrop view of laminated siltstone and lenticular-, wavy-, and nodular-bedded lime
mudstone, with thick bed of burrowed lime mudstone at the top. Scale bar is 5 cm. (B) Negative print of acetate peel showing lenticular and wavy
beds of lime mudstone (dark coloured), containing rare trilobite fragments, and laminated siltstone (light coloured); trilobite grainstone at top.
Scale bar is 1 cm.
Ontario Museum, Toronto (ROM prefix), Geological Survey
of Canada, Ottawa (GSC prefix), United States National
Museum, Washington (USNM prefix), and the Department of
Geology, University of Buenos Aires (UBA prefix).
FAMILY Agnostidae McCoy, 1849
SUBFAMILY Agnostinae McCoy, 1849
GENUS Micragnostus Howell, 1935
Type species
Agnostus calvus Lake, 1906 from the Tremadocian of North
Wales, by original designation.
Remarks
I follow Fortey (1980) (see also Shergold and Sdzuy 1984)
in the concept of this genus. He suggested that the location
of the glabbellar node well behind the transverse glabellar
furrow and the muscle scar pattern are particularly distinctive
characteristics not shared by Geragnostus, with which it has
hitherto been confused. However, illustrations of cephala of
other species of Micragnostus show that many have more ante-
riorly curved transverse glabellar furrows than Fortey (1980,
p. 21) implied and that not all have as tightly "rolled" postero-
lateral borders as Micragnostus serus Fortey, 1980.
Micragnostus intermedius (Palmer, 1968)
(Figs. 5, 8L-8R)
Geragnostus mundus (Raymond, 1925) Lochman, 1964,
p. 464, P1. 63, figs. 39-41.
Geragnostus intermedius Palmer, 1968, p. 24, P1. 12,
figs. 1, 2.
non Geragnostus intermedius, Robison and Pantoja-Alor,
1968, p. 776, P1. 97, figs. 1 - 10.
Geragnostus curvata Robison and Pantoja-Alor, 1968,
p. 775, P1. 97, figs. 11-14.
non Geragnostus (Micragnostus) cf. intermedius, Shergold,
1975, p. 52, P1. 13, figs. 7, 8, 12.
?Geragnostus intermedius, Stitt, 1977, p. 36, P1. 3, fig. 1.
Holotype
A pygidium (USNM 146842) from the Hillard Limestone,
east-centralAlaska, illustrated by Palmer (1968, P1. 12, fig. 2).
Occurrence
Rabbitkettle Formation, Flat River area, District of Mac-
kenzie, Northwest Territories, Ibexian; Deadwood Formation,
east-central Montana (subsurface), Ibexian (Lochman 1964);
Hillard Limestone, east-central Alaska (Palmer 1968), Sun-
waptan; Tiiiu Formation, Oaxaca, Mexico, Tremadocian
(Robison and Pantoja-Alor 1968).
Description
Cephalon is subcircular in outline, approximately as wide as
long, and moderately convex; maximum height is slightly
more than one third the width. Border is wide; marginal fumw
is well incised, wide or wider than rim. Acrolobe is evenly
convex, of the same width around glabella. Glabella occupies
about two thirds the cephalic length and about one third the
 1598 CAN. 1. EARTH SCI. VOL. 25,1988
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FIG. 3 . Negative print of acetate peel showing trilobite-bearing
grainstone overlying lime mudstone. Scale bar is 5 mm.
For personal use only.
FIG.4. Pie diagram showing relative abundance of trilobite genera
in the single collection (N = 196). Unlabelled slice represents
remaining, rare taxa.
cephalic width, measured level with the transglabellar furrow.
Glabella is slightly tapered and evenly rounded anteriorly; it
comprises a circular anterior lobe and an inflated ovate poste-
rior lobe, separated by a transglabellar furrow that forks later-
ally to form a moderately well developed pair of anterolateral
lobes. A faint axial glabellar node is located slightly in front of
mid-length of the posterior glabellar lobe. Basal lobes are tri-
angular and moderately inflated. Genal angles are small blunt
spines.
Thoracic segments are unknown.
Pygidium is subquadrate in outline, approximately as wide
as long, and moderately convex; maximum height is about two
thirds the width. Border is flat; marginal furrow is not well
incised. Unconstricted acrolobe is evenly convex, narrowing
toward rear. Inflated axis is wide and varies from moderately
long to long and parallel sided to slightly inflexed. Anterior
FIG. 5. Micragnostus intemedius (Palmer, 1968). Reconstruction
of cephalon and pygidium. Scale bar is 1 mm.
lateral furrows isolate lateral lobes of first segment. Second
segment is defined posteriorly by a moderately well incised
transverse furrow; it bears an elongate axial pygidial node.
Posterior axial lobe is semicircular; its length is greater by a
third to a half than that of the anterior two segments combined.
A pair of backward-directed posterolateral spines is located
just in front of to just behind the posterior margin of the axis,
depending on its length.
Remarks
Micragnostus intermedius was first described from the Sun-
waptan of Alaska and was defined on the basis of the forward-
curving transglabellar furrow, moderately wide border, and
wide pygidial axis. The Rabbitkettle specimens have more
prominent anterolateral lobes because the posterior fork of the
transglabellar furrow is more incised. Palmer's (1968, p. 24)
original description noted that the glabellar node is situated at
the mid-length of the posterior glabellar lobe. However, the
cephalon illustrated by Palmer (1968, P1. 12, fig. 1) shows the
node anterior of the mid-length, as it is in the Rabbitkettle
specimens. The node is also anteriorly located in Micragnostus
subobesus (Kobayashi, 1936) from the Late Cambrian of
Alaska and Micragnostus haudei Shergold and Sdzuy, 1984
from the Early Ordovician of central Turkey.
Micragnostus cuwata (Robison and Pantoja-Alor, 1968)
from the Tremadocian of Oaxaca, Mexico, is synonymized
here because the supposedly diagnostic forward-curving trans-
glabellar furrow is shared by the type of M. intermedius. On
the other hand, the transglabellar furrow in the Mexican speci-
mens assigned by Robison and Pantoja-Alor (1968) to
M. intermedius is straight, although anterolateral lobes are
faintly present in some. Furthermore, the border furrows are
not as broad, the cephalic border is narrower and more tightly
rolled, and the glabellar node is situated mid-length of the
posterior glabellar lobe. It is therefore probable that these
specimens are not conspecific with M. intermedius. Similar
features of the cephala from the Payntonian of Queensland
described by Shergold (1975) as Geragnostus ( M . ) cf. interme-
dius suggest that they too are not conspecific with the Rabbit-
kettle material. Although the cephala are poorly preserved,
 specimens assigned to G. mundus (Raymond, 1925) by
Lochman (1964, p. 464, P1. 63, figs 39, 41) are almost cer-
tainly conspecific with M. intermedius. As Robison and
Pantoja-Alor (1968, p. 776) noted, this species was based on a
single pygidium and is thus insufficiently known. The cephalon
regarded by Stitt (1977) as G. intermedius seems to have a
slightly forward-curving transglabellar furrow, but it is inade-
quately illustrated, and its synonymy with the Rabbitkettle
specimens is thus uncertain.
The high degree of inflation of the cephalon and pygidium of
M. subobesus, which occurs in the Sunwaptan part of the
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Rabbitkettle in sections farther north (Ludvigsen 1982a), is
considered diagnostic and different from that of M. interme-
dius. The cephalon of M. haudei is similar to the type M. inter-
medius, but the pygidium differs in its posteriorly diverging
margins and tapering posterior axial lobe. Micragnostus
coreanicus Kobayashi, 1960 from the Early Ordovician
Bunkoku Formation of South Korea is also similar, but its
pygidial axis bears a prominent medial ridge rather than a node
and the anterior transverse furrow appears to be complete
across the axis. Most other species of Micragnostus, such as
M. hoeki (Kobayashi, 1939) (see also Jell 1985, P1. 19, figs. 6,
8, 9), have straight transglabellar furrows.
FAMILY Metagnostidae Jaekel, 1909
SUBFAMILY Metagnostinae Jaekel, 1909
GENUS Gymnagnostus Robison and Pantoja-Alor, 1968
For personal use only.
Type species
Gymnagnostus gongros Robison and Pantoja-Alor, 1968
from the Tiiiu Formation of Oaxaca, Mexico, by original
designation.
Remarks
This genus was erected by Robison and Pantoja-Alor (1968,
p. 776) because they regarded the genus Leiagnostus Jaekel,
1909 (type species L. erraticus Jaekel, 1909 from glacial
gravels, northern Germany) as inadequately illustrated and
thus unsuitable for embracing effaced agnostoids of Early
Ordovician age. Even though the holotype of L. erraticus, an
enrolled specimen, has since been reillustrated by Neben and
Krueger (1971, P1. 11, figs. 37, 38), Gymnagnostus still
appears to be a valid genus. The cephalon of L. erraticus lacks
a border, whereas a narrow one is present in both G. gongros
and Gymnagnostus? mexicanus. Pygidia of the holotype and
all other species assigned to Leiagnostus (see, e.g ., Pek 1977)
are characterized by a relatively wide border lacking postero-
lateral spines. These spines are present in G. gongros but not
in G.? mexicanus. Both genera may be related, on the basis of
faint axial impressions preserved on interior molds of some
species of Leiagnostus (e.g., Pek 1977, P1. 4, fig. 5, P1. 5,
figs. 1, 4) and the retention of similar axial furrows in mera-
spids and early holaspids of G. gongros (Robison and Pantoja-
Alor 1968, P1. 97, figs, 28, 33).
Gymnagnostus? mexicanus Robison and Pantoja-Alor, 1968
(Fig. 64)
Gymnagnostus? mexicanus Robison and Pantoja-Alor,
1968, p. 780, P1. 97, figs. 15, 16, 19-22.
Gymnagnostus? mexicanus, Fortey in Fortey et al., 1982,
p. 105, P1. 2, fig. 6.
Holotype
A pygidium (USNM 158883) from the Tiiiu Formation,
Oaxaca, Mexico, illustrated by Robison and Pantoja-Alor
(1968, P1. 97, fig. 22).
Occurrence
Rabbitkettle Formation, Flat River area, District of Mac-
kenzie, Northwest Territories, Ibexian; Tiiiu Formation,
Oaxaca, Mexico, Tremadocian (Robison and Pantoja-Alor
1968); Cow Head Group, western Newfoundland, Tremado-
cian (Fortey in Fortey et al. 1982).
Remarks
A single, tiny pygidium is referred to this species. It is char-
acterized by a narrow border lacking posterolateral spines, a
faint axial node, and very faint anterior portions of the axial
furrows. Pygidia of some species of Leiagnostus, such as
L. bohemicus (NovBk in Perner, 1918) (see Pek 1977, p. 22,
P1. 4, figs. 5 -9, P1. 5, figs. 1-5) are similar, but they have a
more prominent axial node.
FAMILY Leiostegiidae Bradley, 1925
GENUS Leiostegiurn Raymond, 1913
Type species
Bathyurus quadratus Billings, 1860 from the Uvis Con-
glomerate, QuCbec City, Quebec, by original designation.
Remarks
Leiostegium is part of a plexus of closely related taxa charac-
terized by cranidia with short frontal areas and large,
semi-elliptical pygidia with or without a pair of thick marginal
spines. Berg and Ross (1959, p. 113) recognized four sub-
genera of Leiostegium: L. (Leiostegium), L. (Perischodory)
Raymond, 1937, L. (Manitouella) Berg and Ross, 1959, and
L. (Evansaspis) Kobayashi, 1955. Shergold (1975, p. 167)
also followed this scheme. Two additional subgenera were
erected by Zhou and Zhang (1978): L. (Alloleiostegium) and
L. (Euleiostegiurn). Harrington and Leanza (1957, p. 81) and
Lochman-Balk (in Moore 1959) regarded Leiostegium as a
subgenus of Lloydia Vogdes, 1890. However, because the dis-
tinctively swollen pre-occipital part of the glabella of the type
species of Lloydia, L. bituberculatus (Billings, 1865), is not
shared by any other species, Ross (1970, p. 73) considered
Leiostegium a separate genus. I follow Ross (1970) in avoiding
use of the subgenera because of uncertainty whether they
embrace separate, but related, species groups. The various
species can be accommodated under a single genus, Leiosteg-
ium, with the exception of those of Perischodory, as it is
known only from the pygidium that bears a pair of anteriorly
located spines. The pygidium of Evansaspis also bears a pair
of thick spines that extend from the posterolateral margin, but
the cranidium closely resembles that of Leiostegium. Manitou-
ella has wider fixed cheeks and smaller, more anteriorly posi-
tioned eyes than Leiostegium, but the two genera are otherwise
similar. Alloleiostegium is characterized mainly by a border-
less pygidium bearing shallow pleural furrows, although it
may be a junior synonym of Leiostegium, as Jell (1985, p. 55)
implied. Euleiostegium has a rounded anterior margin, but its
resemblance to L. douglasi Hamngton, 1937 suggests it may
also be a junior synonym of Leiostegium. Several species of
Leiostegium have been erected on the basis of pygidia only
(see Shaw 1966, p. 1315); all lack marginal spines.
Leiostegium manitouensis Walcott, 1925
(Figs. 6A-60, 7)
Leiostegium manitouensis Walcott, 1925, p. 104, P1. 23,
figs. 12-19.
Leiostegium manitouense, Ross, 1951, p. 105, P1.27, fig. 1.
Leiostegium manitouensis, Ross, 1957, p. 489, P1. 43, figs.
 1600 CAN. J. EARTH SCI. VOL. 25, 1988

18-20. Hypostome is shield shaped and weakly inflated. Anterior

Leiostegium (Leiostegium) manitouensis, Berg and Ross,
1959, p. 114, P1. 21, figs. 10-13, 17, 24.
Lloydia (Leiostegium) cf. L. manitouensis, Lochman, 1965,
p. 478, P1. 62, figs. 35-41.
Lloydia (Leiostegium) cf. manitouensis, Flower, 1968,
p. 14, P1. 4, figs 3, 4.
~ L ~ d(Leiostegium)
ia sp., Flower, 1968, p. 16, P1. 4, figs.
1. 2. 5.
~eiost&iumsp., Norford et al., 1970, P1. 5, figs. 20, 21.
Leiostegium sp. cf. L. manitouensis, Jell, 1985, p. 64,
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and posterior lobes are not distinctly separated; macula is
shallow. Lateral border is wide and nearly flat, defined by
broad, shallow lateral border furrow. Posterior border is gently
convex, defined by faint anterior border furrow. Anterior wing
is moderately stout..
Pygidium is semi-elliptical in outline and lacks marginal
spines. Gently convex axis tapers backward and reaches poste-
rior border furrow. It is divided into five or six axial rings,
articulating half ring, and terminal piece by transverse ring fur-
rows that are firmly impressed anteriorly but become indistinct

P1. 22, figs. 11, 12.
Diagnosis
A species of Leiostegium with anteriorly rounded, rectangu-
lar glabella. Anterior border short, convex. Palpebral lobe
relatively short, broad, positioned at glabellar mid-length.
Pygidium with moderately wide border defined by shallow
border furrow.
Lectotype
A cranidium (USNM 70350) from the Manitou Formation,
central Colorado, illustrated by Walcott (1925, P1. 23, figs.
12- 14) and designated herein.
Occurrence
Rabbitkettle Formation, Flat River area, District of Mac-
kenzie, Northwest Territories, Ibexian; Manitou Formation,
For personal use only.
posteriorly. Gently inflated pleural field lacks furrows. Lateral
border is wide, gently convex, and defined by faint border
furrow. External surface is smooth.
Remarks
This species exhibits some variability in cranidial shape (see
also Ross 1970, p. 74). Some Rabbitkettle specimens have a
longer, more tapering glabella and longer anterior border, like
those from Tasmania (Jell 1985), and approach the morphol-
ogy of L. douglasi Hamngton, 1937 described from Argentina
(see also Hanington and Leanza 1957, p. 81; Jell 1985, p. 63).
However, the overall combination of short anterior border,
rectangular glabella, and centrally located eyes is distinctive
and separates L. manitouensis from other species.
Leiostegium douglasi has a longer, convex anterior border
and posteriorly located eyes; pygidia appear indistinguishable

central Colorado, Ibexian (Berg and Ross 1959); Jost and
Snake Hills formations, New Mexico, Ibexian (Flower 1968);
Deadwood Formation, east-central Montana (subsurface),
Ibexian (Ross 1957; Lochman 1965); Garden City Formation,
northeastern Utah, Ibexian (Ross 1951); Digger Island Forma-
tion, Victoria, Australia, Tremadocian (Jell 1985).
Description
Cephalon is semicircular in outline and arched. Anteriorly
rounded glabella is moderately convex and subrectangular in
outline in most specimens; rarely it is more elongate and
slightly tapered and only gently convex. It is well defined by
firmly impressed axial furrows that terminate at anterolateral
comers of glabella; glabella may bear three pairs of faint
lateral glabeller furrows. Occipital ring is well defined by deep
occipital furrow. Subtransverse anterior border is very short in
most specimens, is slightly convex, and bears indistinct terrace
lines; rarely it is slightly longer and flatter. Anterior border
furrow is shallow laterally and contains a deep pit at antero-
lateral comer of glabella forward of the anterior termination of
axial furrow. Palpebral lobe is short and broad, located at gla-
bellar mid-length; palpebral furrow is shallow. Anterior trace
of facial suture is straight forward or slightly divergent; fixed
cheek is relatively narrow, about half the width of the glabella,
and may bear a faint, oblique palpebral ridge. Posterior fixed
cheek is moderately narrow and extends laterally a distance
about equal to two thirds the glabellar width. External surface
is smooth.
from those of L. manitouensis. Jell (1985, p. 64) suggested
that Leiostegium Jloodi Shergold, 1975 from the Ninmaroo
Formation of Queensland might be a subjective synonym of
L. manitouensis. However, L. Jloodi possesses a short, flat
anterior border, eyes that are located posteriorly, and a poorly
defined pair of pits in the anterior border furrow. Leiostegium
(L.) planum Peng, 1984 from northwestern Hunan, China,
resembles L. douglasi but differs in having narrow, fixed
cheeks. Leiostegium (L. ) constrictum Peng , 1984, also from
Hunan, resembles Rabbitkettle specimens of L. manitouensis
with elongate glabellas, but its glabella tapers more markedly,
and it has shorter palpebral lobes. Leiostegium mccoyi
(Walcott, 1884) from the Pogonip Group of Eureka, Nevada,
has an anteriorly expanded glabella. Leiostegium puteatum
Raymond, 1924 from the Highgate Formation of Vermont is
based on cranidia similar to that of L. manitouensis but having
posteriorly located eyes. The pygidium assigned by Raymond
(1924, P1. 14, fig. 16) is also similar to those from the Rabbit-
kettle but appears to have a more tapered axis; it was subse-
quently assigned by Shaw (1966, p. 1316) to Leiostegium
elongatum Raymond, 1937, which is characterized by a narrow
border and deeper border furrow. The anterior border of
Leiostegium mudgei Ross, 1958 from upper Ibexian pillow
basalts of Nevada is short and its glabella tapers, and it thus
resembles some specimens from the Rabbitkettle. As Ross
(1970, p. 74) noted, it may be synonymous with Leiostegium
saflordi (Billings, 1860) from the Uvis Conglomerate and

FIG. 6. (A-0) Leiostegiurn manitouensis Walcott, 1925. (A-C) Cranidium: dorsal, anterior, and oblique views, ROM 45748a, ~ 4 .
(D) Cranidium: dorsal view, ROM 49749a, x5. (E) Free cheek: dorso-lateral view, ROM 45750, x6. (F) Cranidium: dorsal view, ROM
. H) Cranidium: dorsal and oblique views, ROM 45752, x 5. (I, J) Latex cast of cranidium: dorsal and oblique views, ROM
4575 1, ~ 4 . 5 (G,
45753a, ~ 3(K, . L) Pygidium: dorsal and oblique views, ROM 45754a, ~ 4 (M,
. N) F'ygidium: dorsal and oblique views, ROM 45755, ~ 3 . 5 .
(0) Hypostome: ventral view, ROM 45756, x 2 . (P) Perischodory incornperturn Berg and Ross, 1959. Fragment of pygidium: dorsal view,
ROM 45753c, x 3.5. (Q) Gymnagnostus? mexicanus Robison and Pantoja-Alor, 1968. F'ygidium: dorsal view, ROM 45754b, X 16.5.
(R) Apatokephalus finalis (Walcott, 1884). Cranidium: dorsal view, ROM 45757, x 13.5. All specimens are from the type area of the Rabbit-
kettle Formation, Flat River, Northwest Territories.
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For personal use only.

PRATT
1601
 1602 CAN. J. EARTH SCI. VOL. 25,1988

obliquely backward-directed, rounded spine at each anterola-

teral comer. Pleural fields smooth. Border moderately narrow.
Remarks
This genus encompasses two species, the type species and
Perischodory incornperturn Berg and Ross, 1959, both known
on the basis of pygidia only. Berg and Ross (1959, p. 113)
argued that Perischodory is a subgenus of Leiostegiurn. How-
ever, in the absence of associated cephala assignable with cer-
tainty to Perischodory and demonstrating a close link with
Leiostegium, I regard it as a separate genus.
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Perischodory incornpertum Berg and Ross, 1959

FIG. 7. Leiostegium manitouensis Walcott, 1925. Reconstruction of
cranidium, free cheek, and pygidium. Scale bar is I cm.
Beekrnantown Formation of southern Quebec. Pygidia of both
are semicircular and have well-impressed border furrows (see
For personal use only.
(Fig. 6P)
Leiostegiurn (Perischodory) incornperturn Berg and Ross,
1959, p. 115, P1. 21, fig. 15, P1. 22, fig. 4.
Lloydia (Perischodory) incornperta, Lochman, 1965, p. 480,
P1. 62, fig. 42.
Holotype
A pygidium (USNM 136421) from the Manitou Formation,
Colorado, illustrated by Berg and Ross (1959, P1. 22, fig. 4).
Occurrence
Rabbitkettle Formation, Flat River area, District of Mac-
kenzie, Northwest Tenitories, Ibexian; Manitou Formation,
central Colorado, Ibexian (Berg and Ross 1959); Deadwood
Formation, east-central Montana (subsurface), Ibexian

also Shaw 1966, P1. 161, figs. 12, 13a, 14, 15), but that of
L. rnudgei has a distinct flat or concave marginal flange, which
is only weakly expressed in L. saffordi. Leiostegiurn tyboensis
Ross, 1970 from the Goodwin Limestone, Nevada, has a
square glabella, a short, highly convex anterior border, and a
very wide pygidial border. Lloydia (Leiostegium) valrnyensis
Lochman, 1966 from upper Ibexian pillow basalts of Nevada
and the upper Ibexian of the subsurface of Montana is a
coarsely tuberculate form with wide and long palpebral lobes.
Two hypostomes from the Rabbitkettle collection, one of
which is figured (Fig. 60), are assigned to L. manitouensis
rather than Kainella eleutherolfi because (i) there is a relative
abundance of the two genera in the collection; (ii) none has yet
been described for Kainella: (iii) one assigned to L. (Evan-
saspis) glabrurn Kobayashi, 1955 (Pl. 2, fig. 16) is similar;
(iv) the transverse anterior margin appears to conform better
with the more transverse anterior border of L. manitouensis
than the arcuate anterior border of Kainella.
GENUS Perischodory Raymond, 1937
Type species
Perischodory grandgei Raymond, 1937 from the Highgate
Formation of Vermont, by original designation (see also Shaw
1966, P1. 161, figs. 3, 4).
Diagnosis
A genus of Leiostegiidae with broad pygidium bearing an
(Lochman 1965).
Remarks
The fragmentary pygidium collected from the Rabbitkettle
shows the spine projecting from the lateral border, just behind
the anterior margin, which is characteristic of P. incornperturn.
In P. grandgei, the spines are extensions of the anterior border.
It is possible that larger collections may show that this subtle
distinction falls within the variability exhibited by one species,
in which case P. grandgei would take priority. The Rabbit-
kettle specimen lacks the shallow border furrow seen in the
types of both these species, but this may be due to tectonic dis-
tortion.
SUPERFAMILY Remopleuridacea Hawle and Corda, 1847
FAMILY Kainellidae Ulrich and Resser, 1930
GENUS Kainella Walcott, 1925
Type species
Hungaia billingsi Wallcot, 1913 from the Chushina Forma-
tion (= lower part of the Survey Peak Formation (Aitken and
Norford 1967, p. 157)) of southeastern British Columbia, by
original designation.
Kainella eleutherolfi n. sp.
(Figs. 8A -8G, 9)
Diagnosis
A species of Kainella with moderately divergent facial

FIG. 8. (A-G) Kainella eleutherolji n. sp. ( A , B) Cranidium: dorsal and oblique views, ROM 45757, ~ 4 (C, . D) Pygidium: dorsal and
oblique views, ROM 45760, ~ 3 . 5 (E)
. Free cheek: dorso-lateral view, ROM 45749b, ~ 3 . 5 (F)
. Pygidium: dorsal view, ROM 45753b, ~ 5 .
(G) Holotype pygidium: dorsal view, ROM 45748b, ~ 3 . 5 (H). Kainella stenorachis Kobayashi, 1953. Holotype pygidium: dorsal view, GSC
11931, x 2, McKay Group, Mount Goodsir, British Columbia. (I-K) Angelina hyeronimi (Kayser, 1876). Cranidium: dorsal and oblique views
and enlargement, ROM 45763, x 5 and x 9.2 (enlargement). (L -R) Micragnostus intermedius (Palmer, 1968). ( L , M) Cephalon: dorsal and
oblique views, ROM 45754c, x 8.8. (N) Cephalon: dorsal view, ROM 45759, x 13.8. ( 0 , P) Pygidium: dorsal and oblique views, ROM 45761,
X8.8. (Q, R) Pygidium: dorsal and oblique views, ROM 45762, x 12.5. All specimens, with the exception of (H), are from the type area of the
Rabbitkettle Formation, Flat River, Northwest Territories.
 Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by University of Saskatchewan on 05/07/14
For personal use only.

'.,..
,'k.
,;
a',
 1604 CAN. J. EARTH SCI. VOL. 25,1988
Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by University of Saskatchewan on 05/07/14
FIG. 9 . Kainella eleutherolfi n. sp. Reconstruction of cranidium,
free cheek, and pygidium. Scale bar is 1 cm.
suture. Lateral glabellar furrows absent. Row of pits located
For personal use only.
forward of anterior border furrow. Pygidium wide, bearing a
single pair of backward-curving furrows and two pairs of
spines.
Holotype
A pygidium (ROM 45748b) from the Rabbitkettle Forma-
tion, Flat River area near Tungsten, District of Mackenzie,
illustrated herein (Fig. 8G).
Name
+
From eleutheros (Greek), meaning free, Rolf Ludvigsen,
my supervisor, in reference to his present state.
Occurrence
Rabbitkettle Formation, Flat River area, District of Mac-
kenzie, Northwest Territories, Ibexian.
Description
Cephalon is semicircular in outline. Slightly convex, anteri-
orly rounded glabella is subrectangular in outline, expanding
imperceptibly at its mid-point and well defined by firmly
impressed axial furrow. Lateral glabellar furrows are absent.
Occipital ring is moderately long and defined by a relatively
short occipital furrow. Preglabellar field is gently down-
sloping, equal in length to slightly convex anterior border, and
separated from it by a moderately long anterior border furrow;
caeca appear to be absent. A row of unevenly spaced pits is
located just forward of anterior border furrow. Crescentric
palpebral lobe is relatively wide and long and equal in length to
half the glabellar length. Anterior branch of facial suture is
moderately divergent at about 20" from transverse; fixed
cheek is narrow. Posterior fixed cheek is very short and is
transverse and extends laterally a distance equal to glabellar
width. Free cheek is broad and gently inflated. Well-impressed
lateral furrow is confluent with posterior border furrow and
defines slightly convex border that extends posteriorly into a
long, flattened spine bearing terrace lines. External surface is
smooth.
Pygidium is subcircular in outline, slightly convex, with
gently tapering, relatively narrow axis composed of six axial
rings, articulating half ring, and terminal piece, which reaches
posterior margin. First pleural furrow is sharply incised and
complete and curves backward; second pleural furrow is very
faint on external surface. Pleural field extends to two pairs of
thick marginal spines that are directed obliquely inward and
terminate beyond the posterior margin.
Remarks
The cranidium of K. eleutherolfi n. sp. differs from all other
species of Kainella in the location of the row of pits in front of
the anterior border. It also differs from that of K. meridionalis
Kobayashi, 1935 from northern Argentina (see Harrington and
Leanza 1957, Figs. 5011 -5013, 5016, 5018) and that of
Kainella billingsi (see also Norford et al. 1970, P1. 5, fig. 31)
in lacking lateral glabellar furrows, having a less divergent
facial suture, and having a noncaecate and slightly shorter pre-
glabellar field relative to the length of the anterior border; the
glabellar of K. billingsi also does not exhibit the slight medial
swelling. The facial suture of Kainella columbianu Harrington
and Kay, 1951 (Pl. 96, figs. 11, 12) from Columbia diverges
at about the same angle as that of K. eleutherolji, but the gla-
bella bears three pairs of firmly impressed lateral glabellar fur-
rows. The elabella of Kainella orentalis Rasetti, 1943 is
u
quadrate rather than subrectangular, and the palpebral lobes
are smaller than those of the Rabbitkettle specimens. Kainella
kindlei Kobayashi, 1955 (Pl. 2, fig. 1) from southeastern
British Columbia differs from K. eleutherolji in having a trans-
verse course of the facial suture, a posterior pair of lateral gla-
bellar furrows, and probably a more convex anterior border.
Pygidia of K. billingsi (see Walcott 1925, P1. 22, fig. 4;
Norford et al. 1970, P1. 5, fig. 32), K. orientalis (see Rasetti
1943, PI. 19, fig. 1I), K. meridionalis (see Harrington and
Leanza 1957, Figs. 5014, 50/5), and Kainella flagricauda
(White, 1874) (see also Lochman 1965, P1. 62, fig. 20) are
characterized by wide axes, narrow pleural fields, and two or
three pairs of furrows directed backward. In contrast, K. eleu-
thorolji possesses wide pleural fields crossed by a single pair
of furrows. The holotype pygidium of Kainella stenorachis
Kobayashi, 1953 (reillustrated in Fig. 8H) from southeastern
British Columbia also has a single pair of pleural furrows, but
the pleural fields are narrow. This pygidium is not laterally dis-
torted, a conclusion confirmed by other taxa from the same
collection (Evans' "Goodsir No. 3717," which is "north of
the first stream from east, north of Brisco trail, elevation
5250 ft") illustrated by Kobayashi (1953, 1955), which are not
deformed. The suture of the cheek assigned to K. stenorachis
by Kobayashi (1953, P1. 3, fig. 6) is more transverse than that
of K. eleutherolji (Fig. 8E) and suggests that this pygidium
and cheek belong to the cranidium of K. kindlei, which is from
the same collection. In that case, K. stenorachis has priority.
The pygidium assigned to K. jlagricauda by Kobayashi (1953,
P1. 3, fig. 5) was found at a different locality in the McKay
Group.
GENUS Apatokephalus Brogger, 1896
Type species
Trilobites serratus Boeck, 1836 from the Tremadocian of
Norway, by original designation.
Apatokephalus finulis (Walcott, 1884)
(Fig. 6R)
 PRATT
Dicellocephalus jnalis Walcott, 1884, p. 89, P1. 12, figs.
12, 12a.
Apatokephalus jnalis, Brogger, 1896, Figs. 6a, 6b.
Diplapatokephalus jnalis, Raymond, 1937, p. 1086.
Apatokephalusjnalis, Lochman, 1965, P1. 62, figs. 3 1, 32.
Cotypes
A cranidium and associated pygidium (USNM 24563) from
the lower Pogonip Group, Eureka District, Nevada, illustrated
by Walcott (1884, P1. 23, figs, 12, 12a) and reillustrated by
Lochman (1965, P1. 62, figs. 31, 32).
Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by University of Saskatchewan on 05/07/14
Occurrences
Rabbitkettle Formation, Flat River area, District of Mac-
kenzie, Northwest Territories, Ibexian; Pogonip Group,
Eureka District, Nevada, Ibexian (Walcott 1884).
Remarks
A single cranidium from the Rabbitkettle type section agrees
in all respects with Walcott's type and hitherto only other
specimen known. Apatokephalus serratus (see Harrington and
Leanza 1957, p. 135) is very similar but has coarsely granular
prosopon and a wider glabellar bulge and thus more arcuate
palpebral lobes. The preglabellar field is absent in other species
of Apatokephalus, including the specimen assigned to A. ser-
ratus by Wilson (1954, p. 275) from an Early Ordovician
boulder in the Woods Hollow Formation of Marathon, Texas.
For personal use only.
FAMILY Olenidae Burmeister, 1843
SUBFAMILY Oleninae Burmeister, 1843
GENUS Angelina Salter, 1859
Type species
Angelina sedgwickii Salter, 1859 from the upper Tremado-
cian of North Wales, by subsequent designation (Vogdes
1890).
Diagnosis
A genus of Oleninae with subtrapezoidal cephalon; glabella
subquadrate, rounded to truncate anteriorly, with lateral gla-
bellar furrows faint or absent. Preglabellar field relatively
long; anterior border furrow pitted. Anterior facial suture
parallel to slightly divergent. Palpebral lobe positioned level
with anterior one third of glabella. Fifteen thoracic segments.
Pygidium relatively small, transverse, with or without short
border spines.
Remarks
This genus has been fully discussed by Henningsmoen
(1957, p. 155), with the correction by Robison and Pantoja-
Alor (1968) to include species with nonspinose pygidia. Ange-
lina spinosa Robison and Pantoja-Alor, 1968 from the
Tremadocian Tiiiu Formation of Oaxaca, Mexico, has well-
developed lateral glabellar furrows, a more divergent anterior
facial suture, a more sinuous trace of the posterior facial
suture, and a more impressed anterior border furrow and has
been assigned to Orygmaspis (Parabolinoides) Frederickson,
1949 by Westrop (1986).
Angelina hyeronimi (Kayser, 1876)
(Fig. 81- 8K)
Arionellus hyeronimi Kayser, 1876, p. 7, P1. 1, fig. 5.
Angelina punctolineata Kobayashi, 1937, p. 479, P1. 6,
fig. 22.
Angelina hyeronimi, Hamngton and Leanza, 1957, p. 99,
1605
fig. 35 (see for synonymy to date).
Angelina punctolineata, Harrington and Leanza, 1957,
p. 101, figs. 3612~-3612e.
Angelina punctolineata, Henningsmoen, 1957, p. 156.
Angelina steinmanni (Kayser, 1897) Henningsmoen, 1957,
p. 157.
Angelina hyeronimi, Robison and Pantoja-Alor, 1968,
p. 785, P1. 101, figs. 1-7.
Angelina hyeronimi, Pnbyl and VanCk, 1980, P1. 8,
figs. 4-6.
Angelina punctolineata, Pfibyl and VanEk, 1980, P1. 8,
figs. 1-3.
Lectotype
A cranidium (UBA 2566) from Tilcuya, Bolivia, illustrated
by Hamngton and Leanza (1957, Fig. 3512).
Occurrence
Rabbitkettle Formation, Flat River area, District of Macken-
zie, Northwest Temtories, Ibexian; southern Bolivia and
northern Argentina, Tremadocian (Harrington and Leanza
1957); Tiiiu Formation, Oaxaca, Mexico, Tremadocian (Robi-
son and Pantoja-Alor 1968).
Remarks
This species was exhaustively described by Hamngton and
Leanza (1957, p. 99), and the single cranidium from the
Rabbitkettle agrees with specimens from South America, as
well as with the Mexican material described by Robison and
Pantoja-Alor (1968). It does, however, possess a more promi-
nently acuminate anterior margin than these specimens, prob-
ably because of postdepositional flattening. Exfoliated
portions of the cranidium exhibit a fine, dense pitting (Fig.
8K), a feature also noted by Robison and Pantoja-Alor (1968).
Angelinapunctolineata was originally based on a single, dis-
torted cranidium from the lower Tremadocian of northern
Argentina. Description and illustrations of additional cranidia
in Harrington and Leanza (1957) do not reveal any characters
different from those of A. hyeronimi. Unless future collections
prove that the pygidia are different, A. punctolineata is
regarded as a subjective synonym of A. hyeronimi. Robison
and Pantoja-Alor (1968, p. 786) also noted that Parabolinella
coelatifrons Harrington and Leanza, 1957 may be conspecific
with A. hyeronomi.
Angelina sedgwicki and Angelina kayseri Harrington and
Leanza, 1957 both have narrower fixed cheeks than A.
hyeronimi. The preglabellar field of A. kayseri is also shorter.
The glabella of A. sedgwicki is longer, and the facial suture is
slightly more divergent. Beltella latifions Wilson, 1954, from
the Lower Ordovician of west Texas, which Henningsmoen
(1957, p. 158) suggested might belong to Angelina, has a deep
anterior border furrow and a quadrate glabella.
No pygidia referrable to A. hyeronimi were collected from
the Rabbitkettle.
Acknowledgments
This paper is a result of fieldwork done as part of doctoral
studies on Cambrian trilobites of the southern Mackenzie
Mountains. I thank Rolf Ludvigsen for supervising this work,
for his companionship on the type section, for his inspiration,
and for critically reading the manuscript. Rube Ross and Brian
Chatterton also made helpful comments on the manuscript as
part of journal reviews. I thank Bill Fritz for his ready advice
 1606 CAN. 1. EARTH SCI. VOL. 25.
on relevant stratigraphic problems. Fieldwork in 1984 was
funded by the Natural Sciences and Engineering Research
Council of Canada and the Department of Indian and Northern
Affairs Canada (northern scientific training grant).
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