Copyright © 1995, The Paleontological Society

SELECTED ORDOVICIAN TRILOBITES FROM THE

LAKE ST. JOHN DISTRICT OF QUEBEC AND
THEIR BEARING ON SYSTEMATICS·
PIERRE J. LESPERANCE AND SYLVAIN DESBIENS
Departement de geologie, Universite de Montreal, P.O. Box 6128, Montreal, Canada, H3C 3J7 and
Geological Survey of Canada, 60 I Booth Street, Ottawa, Canada, KIA OE8

ABsTRAcr- The thorax of Hypodicranotus has ten segments and a spine on the eighth. The ages of Erratencrinurus s.l. spicatus and
Erratencrinurus (Erratencrinurus?) vigilans in the Lake St. John district do not confirm their temporal roles leading to subgenera of
Erratencrinurus, as has been recently suggested. Phylogenetic analyses oflarge data sets of species previously referred to Encrinuroides
and Physemataspis yield a minimal length cladogram containing 18 species. Encrinuroides is restricted to four species, two of which
have biogeographic affinities with Iapetus. These results lead to three clades, named the Walencrinuroides n. gen. clade, Frencrinuroides
n. gen. clade, and finally the Physemataspis clade, with an enlarged concept of the genus with the erection of Physemataspis
(Prophysemataspis) n. subgen. These last three clades are restricted to North America and Scotland, with alternating predominance
of one region. Walencrinuroides s.l. gelaisi n. gen. n. sp. is described. New morphological data on Erratencrinurus s.l. spicatus
confirm its close relationship with the clades discussed above. Data are insufficient for phylogenetic analysis of selected cheirurine
species here surveyed. Eye position, glabellar segmentation, and pygidial shape differentiate the genera Ceraurus and Gabriceraurus;
emended diagnoses of these genera are presented. Ceraurus globulobatus and C. matranseris are distinct, but morphologically close
to one another. The status of Gabriceraurus dentatus can be stabilized on its extant types.

INTRODUCTION Owen and Heath (1990) presented a revision of Erratencrinu-

D ESBIENS ANDLesperance (1989) presented the stratigraphy

of the Lake St. John and Chicoutimi outliers of Quebec
and argued for large-scale northward migration of the faunas
typical of the "middle" Ordovician farther south (Ontario and
New York) into the "upper" Ordovician. They presented sig-
nificant changes in the biozones of selected species, previously
considered diagnostic of specific intervals. The purpose of this
contribution is to document more fully selected remopleuridine,
encrinurine, and cheirurine species and investigate their con-
tribution to evolution and/or systematics.
The age assignments and the arguments for these ages within
the Quebec outliers need not be repeated here. Terminology of
the encrinurines follows Evitt and Tripp (1977).
RECENT CLASSIFICATIONS OF SOME
ORDOVICIAN ENCRINURINAE
The occurrence of three encrinurine species in the Lake St.
John, Chicoutimi, and Manicouagan outliers of Quebec, re-
ported as Encrinuroides vigilans (Hall, 1847), Encrinuroides n.
sp., and Encrinuroides? spicatus (Tripp, 1974) (Desbiens and
Lesperance, 1989), is an opportunity to examine the systematic
bases of some encrinurines. These species were assigned to the
Edenian, the basal stage of the Cincinnatian Series of North
America, which underlies the Maysvillian Stage. The Carado-
cian-Ashgillian boundary is now known to be at, or very near,
the EdeQian-Maysvillian boundary (Barnes et aI., 1981; Berg-
strom and Mitchell, 1986). The reported ages of the three taxa
quoted above, however, do not accord with the evolutionary
paths recently suggested. Therefore, these recent suggestions are
briefly reviewed below.
The genus Erratencrinurus Krueger, 1971. -Encrinurus spi-
catus has acquired pivotal significance in the evolution of some
"upper" Ordovician encrinurines through the numerical anal-
yses of Temple and Tripp (1979). They recognized this species
as leading to two distinct branches, later formally recognized as
Erratencrinurus s.s. and Erratencrinurus (Celtencrinurus) Evitt
and Tripp, 1977 (Strusz, 1980; Lesperance and Tripp, 1985).
1 GSC contribution no. 11994.
rUSe They suggested a phylogeny of all the species, without un-
dertaking a rigorous cladistic or stratophenetic analysis due to
deficient stratigraphical and systematic knowledge of some spe-
cies (Owen and Heath, 1990, p. 228). They recognized two
species groups within Erratencrinurus (Erratencrinurus), two
additional ones within Erratencrinurus (Celtencrinurus), and an
Erratencrinurus s.l. group; they also presented subgeneric di-
agnoses, but deferred a generic diagnosis until generic diagnoses
of other encrinurine genera were finalized. Erratencrinurus s.1.
spicatus was considered a morphological and temporal inter-
mediate between all Erratencrinurus species and the probable
ancestral lineage from Encrinuroides torulatus Evitt and Tripp,
1977, to Encrinuroides uncatus Evitt and Tripp, 1977. Erra-
tencrinurus (Erratencrinurus?) vigilans [sic] was considered in-
termediate in position between spicatus and Erratencrinurus
(Erratencrinurus) species.
Both Strusz (1980) and Owen and Heath (1990) assigned a
mid-Caradocian age to spicatus, then only known from the Green
Bay area of Wisconsin, and reported as originating from the
Galena Formation (Tripp, 1974, p. 484). Recent correlation
charts recognize a dozen or so members within the Galena, and
the lack of precise stratigraphic information in Tripp (1974)
renders its exact age problematical. The Galena of southeast
Minnesota, some 500 km westward of the Green Bay area, has
been assigned to the tenuis and conjluens conodont zones (Sher-
manian-Edenian; Sweet, 1984) and thus is of mid- to latest
Caradocian age (Barnes et aI., 1981). Available evidence on the
age of spicatus, both from its type area and the Lac Saint-Jean
and Manicouagan outliers (where it is latest Caradocian), con-
sequently does not support chronostratigraphically its ancestral
relationship with all the species of Erratencrinurus. New mor-
phological data on spicatus presented below confirm its close
relationship with species ofthe widely conceived genus Encrinu-
roides Reed, 1931.
It appears reasonable to accept both Ludvigsen's (1979a, p.
45) and DeMott's (1987, p. 80) judgments that Encrinurus tren-
tonensis Walcott, 1877, and E. cybeleformis Raymond, 1921,
are junior synonyms of "Encrinurus" vigilans (Hall, 1847), as
the specific concepts of the first two are based on pygidia. Thus
conceived, vigilans extends from the Rocklandian to the Eden-
ian of southern Ontario (Ludvigsen, 1978) and also occurs in

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 2 LESPERANCE AND DESBIENS
the Edenian of the Manicouagan outlier (Desbiens and Lespe-
rance, 1989); that is, it occurs from early Caradocian (Harna-
gian) to latest Caradocian (Barnes et aI., 1981). This again alters
significantly the stratigraphic relationships recognized by Owen
and Heath (1990). The Manicouagan outlier material ofvigilans
does not furnish new data on this species; the lectotype chosen
by DeMott (1987) has been illustrated by Ludvigsen (1979a, fig.
27A) and Edgecombe and Chatterton (1990, PI. 1, fig. 11).
Owen and Heath's (1990) Erratencrinurus s.1. sp. B [=Erra-
tencrinurus (s.1.) of Lesperance and Tripp, 1985] occurs in the
Rouge Member of the Pabos Formation, assigned to the Pus-
gillian-Cautleyan (Ashgillian) (Lesperance et aI., 1987) and not
to the late Caradocian as given by Owen and Heath (1990, fig.
2). This change of age does not alter Owen and Heath's (1990)
interpretation of evolution within their Erratencrinurus s.l.
branch.
Edgecombe and Chatterton (1990) examined critically some
generic concepts within the Encrinurinae using cladistic meth-
ods. They accepted that Erratencrinurus formed a monophyletic
group, noted that Encrinuroides was in need of revision, and
that possibly many new genera may be required to assign all
species to monophyletic taxa (Edgecombe and Chatterton, 1990,
p. 823). They considered that, as presently conceived, Encrinu-
roides was paraphyletic and they discussed various recent di-
agnoses of the genus. The majority of the characters used re-
cently were considered as synapomorphies of the encrinurines
or as plesiomorphies shared with other groups. They also point-
ed out that some species were assigned to Encrinuroides on the
absence of apomorphies with other genera.
Temple and Wu (1990) amplified the work of Temple and
Tripp (1979) by the addition of numerous Chinese species. Or-
dination, including the Chinese species, isolated a fairly tight
group of "Encrinuroides-like forms" (Temple and Wu, 1990, p.
21 7), indicating survival of the morph in the Silurian of China
after its disappearance elsewhere (p. 218). Silurian encrinurines
are consequently excluded from this contribution.
The genus Physemataspis Evitt and Tripp, 1977. -Physe-
mataspis mirabilis Tripp, 1980a, was added to the monotypic
Physemataspis coopi Evitt and Tripp, 1977, by Tripp (1980a).
Edgecombe and Chatterton (1990, p. 823) suggested that En-
crinuroides insularis Shaw, 1968, was more informatively clas-
sified in Physemataspis. Furthermore, following Strusz (1980),
they suggested that the Llandeilian-Caradocian group composed
of Encrinuroides periops Tripp, 1967, E. tholus Evitt and Tripp,
1977, E. capitonis Frederickson, 1964, E. pernodosus (Slocom,
1913), Encrinuroides sp. of Ross and Shaw, 1972, with possibly
E. gibber (Dean, 1979) and two poorly known Late Ordovician
species, was more closely related to Physemataspis than to a
restricted monophyletic concept of Encrinuroides. A new species
(reported above as Encrinuroides n. sp., and described below
as Walencrinuroides s.l. gelaisi n. gen. n. sp.) is more easily
compared with North American species than with other known
ones.
PHYLOGENETIC ANALYSIS OF ENCRINUROIDES SPECIES
During the review process for this contribution, it was strongly
suggested that the traditional systematics used for selected En-
crinurinae be abandoned and that cladistic parsimony analysis
be employed. This is certainly not the place to review these two
opposing philosophies and methods; phylogenetic methods are
described in recent contributions by Wiley et aI., 1991, Forey
et aI., 1992, and Smith, 1994. It is nonetheless pertinent to note
that reversal of character states in cladograms is common, a
situation which in traditional systematics is dealt with by rel-
egating these reversals to a lower taxonomic level, or ignoring
them altogether as undiagnostic. The choice of characters and
included taxa, however, merit extensive discussion.
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Recent analyses of encrinurines by Temple and Tripp, 1979,
Temple and Wu, 1990, and Edgecombe and Chatterton, 1990,
have provided numerous characters and basic raw data to use.
Without the first two studies, it would have been impossible to
assemble a reasonable matrix of characters. Nonetheless, the
choice of characters is delicate and somewhat subjective. It is
commonly stated that as many characters as possible should be
assembled for any analysis (e.g., Smith, 1994, p. 35, 43, 45).
This approach may be fine in theory, and even desirable and
necessary, but even modern computers place a limit on the size
of the matrices that can be analyzed (discussed more fully be-
low). In addition, knowledge of the taxa under study varies
widely according to the nature of the material and its complete-
ness of description. Biological interpretations of many of these
characters is at best difficult, especially in extinct groups such
as the trilobites, and one has the feeling of dealing with totally
abstract concepts of taxa of unknown nature.
Following Edgecombe and Chatterton's (1990) review, phy-
logenetic analysis of taxa included within Physemataspis and
the broadly conceived genus Encrinuroides was judged worthy
of investigation. Taxa included in Erratencrinurus, as outlined
by Owen and Heath (1990), including Erratencrinurus s.l. spi-
catus and E. (Erratencrinurus?) vigilans, were excluded from
consideration, as previous investigators have accepted this as a
monophyletic group. The genus Physemataspis, obviously re-
lated to a broadly conceived Encrinuroides through insularis
Shaw, 1968, appears to be the sister group of Encrinuroides,
and was considered as such in the early stages of analysis. A
survey of most species conceivably members of Encrinuroides,
gleaned from original or later descriptions, Temple and Tripp
(1979) and Temple and Wu (1990), isolated 18 taxa well enough
illustrated, described, or preserved for consideration (Table 1).
Taxa not in this table, possibly or probably best included in
Encrinuroides, are either based on pygidia or do not meet min-
imal requirements for determination of character states.
Characters considered and retained. -Smith (1994) has dis-
cussed at length the qualities that characters used in phylogenetic
analysis should possess. Of primary importance in the choice
of these characters is the objectivity of their definition and the
assignment to their separate states. Particular difficulties are
associated with continuous variables. Stevens (1991) has out-
lined the problem and, as a consequence, the writers have at-
tempted to eliminate continuous variables. Many of Temple
and Tripp's (1979) variables are continuous, and between six
and nine ofthe 28 variables ofEdgecombe and Chatterton (1990)
are continuous. Nonetheless, some characters are inherently
variable and as a consequence Encrinuroides hornei Dean, 1973,
the oldest of the encrinurines (latest Arenigian or earliest Llan-
virnian), was used to define character states 0 and GSC primary
types 32743-32749 were examined to complement the pub-
lished description. Assignment of character states 0 to hornei is
an attempt at reducing the subjectiveness of determining char-
acter polarities. Determining polarity of characters with strati-
graphic data is fraught with difficulties (see discussion, chapter
3 in Smith, 1994), but the procedure was adopted here for lack
of other alternatives. Not all characters retained here can be
observed in hornei and as a consequence the earliest cybelines
have been used to determine some character states.
The cybelines are probably the sister group of encrinurines,
as implied by Edgecombe and Chatterton (1990); they share
forwardly expanding glabellae, median glabellar furrows on the
frontal lobe, and pauci-segmented pygidia with similar archi-
tecture. The cybeline Lyrapyge ebriosus Fortey, 1980, is older
than the earliest encrinurine (hornei), and is possibly the oldest
cybeline (Fortey, 1980, p. 100); the slightly younger cybelines
which are probably also older than hornei, Cybelurus brutoni
 ORDOVICIAN TRILOBITES 3

Fortey, 1980, succeeded by C. halo Fortey, 1980, were also used. TABLE 1- Taxa retained for cladistical analysis; last three taxa are cybe-
Character states 0 were determined from these early cybelines lines used for determination of ancestral states (see text for further
details).
when they could not be observed in hornei (notably thoracic
and hypostomal data).
Author
A significant number ofcharacters were not coded for a variety Taxon (revision or redescription)
of reasons. Two characters are constant within the 18 taxa re-
Encrinuroides autochthon Tripp, 1962
tained: minimal glabellar width occurs across the L 1 lobes and Encrinuroides capitonis Frederickson, 1964 (Shaw, 1974)
there is absence ofa sagittal depression on the anterior cranidial Physemataspis coopi Evitt and Tripp, 1977
border (its presence is diagnostic [apomorphic] of Erratencrin- Walencrinuroides s.l. gelaisi herein
urus). These two plesiomorphic characters are consequently not n. gen. n. sp.
instructive. Four other characters are too subjective for use. The Encrinurus gibber Dean, 1979 (Edgecombe and Chatter-
ton, 1990)
number of glabellar tubercles and the tubercle size index (Tem- Encrinuroides hornei Dean, 1973 (herein)
ple and Tripp, 1979) are continuous variables and there is no Encrinuroides insularis Shaw, 1968
obvious way of assigning states to these measures. The sym- Encrinuroides lapworthi Tripp, 1980b
metrical arrangement of the anterior border tubercles is seldom Encrinuroides neuter Evitt and Tripp, 1977
Encrinuroides obesus Tripp, 1965
well described, or illustrated, and subjective in assessment. The Encrinuroides periops Tripp, 1967
absence or presence of the torular tubercle (Evitt and Tripp, E ncrinuroides polypleura Tripp, 1967
1977) is difficult to determine from illustrations, is seldom de- Ceraurus rarus Walcott, 1877 (Chatterton and Lud-
scribed (as it has been recently defined) and, furthermore, occurs vigsen, 1976; DeMott, 1987)
Cybele sexcostatus Salter, 1848 (Whittington, 1950,
in some species only during their ontogeny. Finally, the absence 1965; Price, 1974)
or presence of glabellar tubercle pairs requires either excellent Encrinurus stincharensis Reed, 1928 (Tripp, 1979)
descriptions or well oriented illustrations for accurate deter- Encrinuroides tholus Evitt and Tripp, 1977
Encrinuroides torulatus Evitt and Tripp, 1977
mination. Encrinuroides uncatus Evitt and Tripp, 1977
Seventeen characters were retained in the phylogenetic anal- Lyrapyge ebriosus Fortey, 1980
ysis. These are described and commented upon in the Appendix. Cybelurus brutoni Fortey, 1980
Character polarity. - As previously alluded to, this subject is Cybelurus halo Fortey, 1980
fraught with difficulties, which can lead to tortuous assumptions
and reasonings. In an effort to bypass these difficulties, realizing
that the group under investigation has always been considered Data from Tables 1 and 2 were thus subjected to cladistic
as the early radiation of the encrinurines, character states 0 parsimony analysis, as discussed above. Figure 1 gives the unique
common to the earliest encrinurine (hornei) and the earliest solution from the data in the tables; this cladogram was obtained
cybeline (Lyrapyge ebriosus) were defined as Ancestral State 1 through branch-and-bound search. Three well individualized
ofTable 2. Ancestral State 2 are those character states 0 common clades issue from the basal nodes of the cladogram and are
between hornei and Cybelurus brutoni (ifabsent from C. brutoni, assigned generic status. It is gratifying to note that the cladogram
these states were taken from the slightly younger C. halo). It is retains some previously described relationships between some
submitted that under the circumstances this method is the least species, as autochthon-polypleura (Tripp, 1967), coopi-insularis
subjective method that can be suggested. (Edgecombe and Chatterton, 1990), and neuter-uncatus (Evitt
Results of analyses. - Because of the large matrix at hand and Tripp, 1977; Edgecombe and Chatterton, 1990).
(Table 2), PAUP 3.1.1 installed on a Power Macintosh 6100/ Figure 1 reveals a close relationship between insularis, coopi,
60, with 256 Kb cache memory, was chosen as the most efficient neuter, and uncatus. This clade also forms a monophyletic group
combination for cladistic analysis. Searches for the shortest trees ifconsidered as the outgroup to the other species, in cladograms
were performed using different strategies. The practical limit of
exhaustive searches for all trees is, approximately, a data matrix
of 12 taxa and 14 characters (which evaluated 6.5 x 108 trees
in an interval of 36.5 hours). Heuristic (approximate) searches TABLE 2 - Data matrix of the 18 taxa retained for cladistical analyses,
with two ancestral states, as discussed in the text.
are useful, but the general approach yields spurious results based
on order of data entry, which unfortunately occurred during the
Character states
analyses. Branch-and-bound searches, with furthest addition se-
quence, are thus indicated, even if less ideal than exhaustive Taxon 1 234 5 6 789 10 11 12 13 14 15 16 17
searches. Available computer hardware does not permit ex- autochthon 01011 1 100 1 1 1 ? 0 0 0 1
haustive searches of data matrices other than relatively small capitonis o10 1 1 1 1 10 0 1 1 1 0 0 1 0
ones· even branch-and-bound searches as detailed above have coopi 100 1 1 1 1 1 2 0 0 1 1 1 0 1 0
gelaisi n. sp. 000 1 1 1 102 0 1 1 1 ? 0 ? 1
an i~precise upper limit of approximately 20 taxa. If only a gibber 010 I I ? 1 ? 2 0 1 1 ? ? 0 ? 0
restricted set can be effectively analyzed, the steps leading to hornei o0 0 0 0 0 0 0 0 0 0 0 ? ? 0 ? 0
exclusion of taxa are subjective, which goes against all the aims insularis 000 1 1 1 112 0 0 1 ? 0 0 1 0
of objectivity of phylogenetic analysis. lapworthi 000 1 1 1 1 1 1 0 1 0 1 0 0 0 1
neuter 000 1 1 1 1 1 1 0 0 0 0 1 0 0 0
The first attempts isolated the data structure using Physe- obesus o 0 0 0 1 0 100 0 1 1 ? 0 0 0 0
mataspis coopi as the outgroup; insularis was always its closest periops 12011 1 102 1 1 1 ? 0 0 0 0
associate and tree lengths of 36-38 were commonly obtained, polypleura 010 1 1 1 100 1 ? 1 ? 0 0 0 1
with matrices of the same size as Table 2, but differing in a few rarus 010 1 1 1 101 0 1 1 1 0 1 1 1
characters. Ancestral State 1 was employed in a second step to
sexcostatus o 0 101 1 100 0 1 2 1 ? 0 ? 0
stincharensis 000 1 1 1 102 0 1 1 ? 0 0 0 1
polarize the characters. Ancestral State 2 was also used to po- tholus 010 1 1 1 1 1 2 0 1 1 ? ? 0 ? 0
larize the characters and it gave identical results to those using torulatus 010 1 1 1 1 1 0 1 1 1 1 0 0 1 0
Ancestral State 1, except that the retention index obtained was uncatus 000 1 1 1 1 1 0 0 0 0 0 1 0 1 0
Ancestral state 1 0 000 0 0
lower (0.714 vs. 0.731); this ancestral state is consequently not Ancestral state 2 0 o0 0 0
considered any further here.

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 4 LESPERANCE AND DESBIENS

I I I I
I I I I
I I I I

: Encrinuroides : Walencri nuroides Frencrinuroides : Physemataspis :

I I I I
I I I I

~
I
J ~~c:, ~
I
J ~ti,'\.
~
~ ~ ~J ~'CD
I ®~ ®II
~~~.~ ~c:, r:"fli. nc:, ~-:~.~ ~0 ~~\ ~~O 0~(. ~~ ~\c:, ~~c:, ~ ~ ~\c:, 'it.~c:, ~
~ ~ ~~0 0'5 ;:f..CO ~\Oy~ ~ ~~Cj • C~ ~O '!. ~c:, ~Ov ~\~~~ ~~~0'( O~~c:, .n~O <I~fli. ~ ~ ~~fli. O-Q' ~,0 ",C'l;.~ ~~
~O O~ c:,0 ~0 .," ~0 c:,~~ \'l>'Q '(fli. 'l>~ ~O\1 ~ ~v ~ c~Y ,0' ~ ~ \~~ CO ~0 ~'" ~ ~

(16,1)
I
(15,1)
U (10,1)
I
(9,0) (9,0)
I
(10,1)

(1,1)
(1~.O)
(9,1)

(12,0) (13,0)
(9,0)

(12,0)
I
(8,1)
I (2,1)
I (14,1)

(10,1)
I (9,1) (2,1)
I
(11,0)
I
I I
(2,2) (16,1 )
I I
(1,1) (17,1) (8,1)
I I I
I
(12,2) (9,2)
I I
(3,1) (4,1)
I I

(6,1)
I
I

(7, 1) (12, 1)
(5,1) (11,1) CD Physemataspis (Physemataspis)
I ® Physemataspis (Prophysemataspis)
(13,1)
I
FIGURE 1-Unique minimal length cladogram of the 18 encrinurine taxa of Table 2, using the branch-and-bound method, with furthest addition
sequence, of PAUP and Ancestral State 1 of Table 2. Tree length is 34, consistency index 0.588, and retention index 0.731. Changes from
initial character states 0, as well as reversals, are indicated above their respective nodes (first number is the character and the second its state).

not presented here. As coopi is the type species of Physemataspis, oftype species, gibber and tholus are assigned a sensu lato status.
its generic concept is best broadened to include neuter and un- The remaining taxa of Figure 1 could be referred to Encrinu-
catus, which by themselves form a compact subgroup isolated roides. However, the gelaisi-stincharensis-lapworthi-rarus-au-
at the subgeneric level as P. (Prophysemataspis) n. subgen. A tochthon-polypleura clade is obvious. This clade can be easily
sensu lato assignment of insularis to Physemataspis is conse- defined and is herein assigned to Walencrinuroides n. gen., with
quently indicated. rarus as type species, necessitating a sensu lato assignment to
Physemataspis (Prophysemataspis) n. subgen., as defined four species.
herein, has been assigned an important role, at least, or has been It follows, from the assignment of character states 0 to hornei
considered the root stock for younger encrinurine genera by and the method of defining the ancestral states, that hornei
numerous investigators (Evitt and Tripp, 1977; Strusz, 1980; would be the closest taxon to the hypothetical ancestor. It ap-
Edgecombe et aI., 1988; Owen and Heath, 1990; Edgecombe pears logical that Encrinuroides be restricted to the clade con-
and Chatterton, 1990). This subject is, however, outside the taining hornei to periops, although hornei and obesus must be
scope of this contribution. assigned a sensu lato status within Encrinuroides.
Another well individualized clade that stands out from the The choice of Encrinuroides capitonis Frederickson, 1964,
parsimony analysis is the gibber-tholus-capitonis-torulatus and Ceraurus rarus Walcott, 1877, as type species of Frencrinu-
group. This clade is herein named Frencrinuroides n. gen., with roides n. gen. and Walencrinuroides n. gen., respectively, is dic-
capitonis as type species. As shown in the cladogram, this is a tated by the necessity of using the best known species within
morphologically intermediate clade, and because of the choice each clade. This has the unfortunate consequence of assigning

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 ORDOVICIAN TRILOBITES
the phylogenetically previous species of each clade a sensu lato
status.
Biogeographic and age considerations. -Support of the clado-
gram of Figure 1 can be gathered from the spatial and temporal
distributions of the various clades. Encrinuroides, as restricted,
has biogeographic affinities with Iapetus (hornei in eastern New-
foundland and sexcostatus in northern Wales), while the other
included species occur in Scotland. The duration of this group
is, however, the most extensive, as it occurs from the Arenigian-
Llanvirnian to the Ashgillian. Walencrinuroides n. gen. is main-
lya Llanvirnian-Llandeilian clade, extending to the middle (lap-
worthi) and upper (gelaisi n. sp.) Caradocian. It occurs mainly
in Scotland and less so in cratonic North America. The Fren-
crinuroides n. gen. clade is retricted to eastern cratonic North
America and the adjacent mioclinal rocks of Virginia; it is re-
stricted to the lower and middle Caradocian. The Physemataspis
clade is again a lower and middle Caradocian clade, with the
exception of insularis, which is slightly older (Llandeilian), cor-
responding to its position in the cladogram. This Physemataspis
clade has the same spatial distribution as the Frencrinuroides
n. gen. clade, with the addition of Scotland (Physemataspis mi-
rabilis), a region well documented for its affinities with Virginia
trilobites of the same age. As a whole then, these observations
reinforce the cladogram of Figure 1.
Concluding remarks. -Species based on pygidia, and with
unknown cephala, are difficult to assign unequivocally and bet-
ter referred to "Encrinurus" until the phylogeny of the Encrinu-
rinae is more fully understood.
Additional described material or illustrations are sorely need-
ed for elucidating the status of the following species: Encrinurus
contentus Reed, 1914 (PI. 6, figs. 11, 12), to which Tripp, 1962,
p. 24, referred the cephalon illustrated as Cybele? perversa Reed,
1935 (PI. 4, fig. 5; also, Tripp, 1967, p. 71); Encrinurusfallax
Reed, 1899 (p. 753, PI. 49, figs. 9-12; for comments, see Tripp,
1962, p. 24, who described E. autochthon's cranidial construc-
tion are similar to that of E. fallax); Encrinurus pernodosus
Slocom, 1913 (p. 61, PI. 16, figs. 5-7); E ncrinuroides waigatch-
ensis Burskyi (1966, p. 79, figs. 5-15); and Encrinuroides sp. of
Westrop and Ludvigsen, 1983 (p. 23, PI. 8, figs. 11, 12, 14).
Until this is done, these species and others not discussed here
are best assigned to Encrinuroides?
THE CHEIRURINE GENUS CERAURUS AND ITS ALLIES
The number of species that have been assigned to Ceraurus
Green, 1832, is impressive, as it is a long standing genus, its
putative species are widespread, and probably restricted to North
America in addition to being long ranging ("middle to upper"
Ordovician). As have all genera that were suggested in the early
nineteenth century, it has been frequently dismembered through
the recognition of distinct genera with diagnoses that are not
always distinctive.
A phylogenetic study of the taxa centered on the genus Cerau-
rus has not been attempted because of the absence of a large
data base, as available for the encrinurines. Ludvigsen (1979b)
remains the most important contribution to our knowledge of
the genus to date; this contribution complements Ludvigsen's
(1979b) important survey. As with many older species, many
cheirurine species remain incompletely known and, most im-
portantly, their intraspecific variation is seldom detailed, ana-
lyzed, or even mentioned. As for the genus Cryptolithus (Shaw
and Lesperance, 1994), significant intraspecific variation is sus-
pected (and some is detailed below). A data matrix for phylo-
genetic analysis ofthe 25-30 species discussed here would prob-
ably be unresolvable with present computing means. A tradi-
tional treatment of the systematics of the few cheirurine genera
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5
investigated here appears the only course of action presently
available.
The type specimen of Ceraurus, pleurexanthemus Green, 1832,
is totally unsatisfactory in preservation (Evitt, 1953, PI. 5, fig.
1), and knowledge of the species comes from other material
superbly illustrated by Evitt, 1953, from the same general area
west of the Adirondacks. Characters that are commonly used
for generic diagnoses in other cheirurids or other trilobite fam-
ilies show an apparent haphazard mixture in distinct species
and populations. Thus, diagnostic generic characters are not
obvious within Ceraurus and its closest allies with forwardly
expanding glabellae, the genera Gabriceraurus Pribyl and Vanek,
1985, and Leviceraurus Hessin, 1988a. These three genera share
similar hypostomata, stout, more or less elongated genal spines,
non-carinated large pygidial spines originating from the anterior
pygidial axial ring, an ornamentation of a mixture of pustules
and/or tubercles, and isolated, or at least circumscribed, Ll
lobes. The importance ofexoskeletal convexity is problematical,
and it is suspected that it is related to size and hence not properly
a character that can be used at the generic level. (Extensive
collections of Ceraurus pleurexanthemus from the St. Lawrence
Lowlands of Quebec show that collections with individuals of
greater average size than other collections have greater convexity
and more frontal glabellar protrusion.)
Most of the North American cheirurine species near pleurex-
anthemus can be assigned confidently to Ceraurus and Gabri-
ceraurus, following their emended diagnoses presented below.
The genus Leviceraurus is perhaps a valid taxon, but will not
be discussed further, as it only includes its type species Levi-
ceraurus mammilloides Hessin, 1988a.
The genus Gabriceraurus, erected by Pribyl and Vanek, 1985,
with Ceraurus gabrielsi Ludvigsen, 1979b, as type species, was
accompanied by an uninstructive diagnosis. This new genus was
subsequently used without comment by Shaw and Tripp (in
DeMott, 1987) and by Hessin (1989). The obvious distinction
between Gabriceraurus and Ceraurus lies in the relative position
(exsag.) of the eyes, farther backward in the former (opposite
L2) than in the latter (L3). Even this character is not entirely
constant in natural populations and between the species assigned
below to the two genera. Previous investigators had assigned to
Ceraurus species with eyes in both positions, not recognizing
this criterion as of generic significance. Nonetheless, careful at-
tention to glabellar segmentation, inflation of the anterior gla-
bellar lobe, and the nature of the posterior pygidial margin
between the great spines provides criteria complementary to eye
positions. "Tendencies" are considered morphologic traits more
commonly found in one taxon rather than in the other, but
which are not restricted to either taxon. Even with revised di-
agnoses, some species remain of uncertain affinities (see below).
Both Ceraurus and Gabriceraurus are useful taxa, and they
are formally and more fully diagnosed (and emended) in the
Systematic Paleontology section. Ceraurus has eyes in an an-
terior position, in conjunction with a glabella that either over-
hangs or nearly reaches the frontal margin, a glabellar segmen-
tation that is restricted laterally, and a posterior pygidial margin
that tends to be subtriangular. In contrast Gabriceraurus has a
frontal border that is always visible when viewed dorsally, in
conjunction with eyes that are set more backward, and a glabella
that is more segmented proximally than that of Ceraurus; its
posterior pygidial margin between the great spines tends to be
transverse, with a tendency to spinosity. These data are more
evident on Figure 2, which shows graphically these distinct gla-
bellar and pygidial outlines. Glabellar ornamentation in Cerau-
rus tends to be symmetrically disposed, while in Gabriceraurus
glabellar ornamentation is seldom symmetrical.
As thus conceived, Ceraurus includes C. cetus Dean, 1979,
 6 LESPERANCE AND DESBIENS

Ceraurus

Pleurexanthe~:~EN +-0-
18
-rt-
2Y
-0- 3~
~4
~
0 "--/ ~
pleurexanthemus
montyensis EVITT
+-(O~t
~
- _rR()---.
~\ R ~9 >g.
f/; ~

whittingtoni EVITT :Q~Q~~Q ~

globulobatus
BRADLEY 180 190- ~O-
-0 -0- 0-
C-..=:> c=s c::::=:s
9IObUIOo~a~~~SIN
6
cf.

26\:1- 27(/ :y-

22b 23~ 24

n)---.
matranseris
SINCLAIR

milleranus
MILLER & GURLEY 31
0---.
c::::5
GD~ A
C=s13---. +-
32
~
33 ~

milleranus
of LUDVIGSEN
350-
ruidus
COOPER 3:0- ~fjj- :Q-
Gabriceraurus

gabrielsi
LUDVIGSEN ~
~
blussoni ~
LUDVIGSEN
~
hirsuitus
LUDVIGSEN

dentatus
RAYMOND & BARTON
59

plattinensis
FOERSTE

FIGURE 2 -Outline drawings of various species of Ceraurus and Gabriceraurus showing their respective glabellae, occipital rings, and pygidia.
Transverse arrows point to the mid-point (exsag.) of the palpebral lobes when known. Proximal limits of glabellar furrows shown by small
filled circles, joined together by imaginary lines. Source of drawings are as follows. 1, Evitt, 1953, PI. 5, fig. 5 x 1.4; 2-4, Whittington, 1941,
PI. 73, fig. 4 x 1.3, 3 x 2.7, 24 x 4.0; 5, 6, Evitt, 1953, PI. 8, fig. 5 x 1.8, 6 x 3.1; 7-11, Evitt 1953, PI. 5, fig. 9 x 1.4, 11 x 2.2, PI. 8, fig. 12
x4.6, 11 x4.5, 8 x 1.9; 12, 15, Whittington, 1992, PI. 2, fig. A x 1.4, PI. 3, fig. A x2.8; 13, 14, 16, 17, Evitt, 1953, PI. 5, fig. 16 x2.3, 14
x 1.5, PI. 8, fig. 25 x3.1, 23 x 1.9; 18-21, herein, Fig. 4.14 x 1.7,4.16 x 1.7,4.17 x 1.3,4.22 x 1.8; 22-25, Hessin, 1989, PI. 4, fig. 1 xLI, 4
x 1.7,5 x 1.6,3 x 1.6; 26-30, herein, Fig. 4.9 x4.1, 4.10 x 1.2,4.3 x 1.8,4.12 x 1.8,4.13 x 1.5; 31-34, Raymond and Barton, 1913, PI. 1,
fig. 8 xLI, 6 x 1.9,7 x 1.6, PI. 2, fig. 6 xO.9; 35,36, Ludvigsen, 1979b, PI. 15, fig. 46 x2.2, 53 x2.8; 37-40, Shaw, 1974, PI. 7, fig. 1 x 1.4,
15 x 1.7, 10 x 3.3, 5 x 2.1; 41-46, Ludvigsen, 1979b, PI. 12, fig. 1 x 1.1, PI. 11, fig. 1 x 1.1, PI. 12, fig. 4 x 1.4, 14 x 1.3, 25 x 2.2, 18 x 3.3;

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 ORDOVICIAN TRILOBITES
PI. 2, figs. 1-5, 9, PI. 4, fig. 10; C. globulobatus Bradley, 1930,
C. cf. C. globulobatus of Hessin, 1989, non Bradley, 1930, and
C. matranseris Sinclair, 1947, all three of which are discussed
at some length in the Systematic Paleontology section; C. mil-
leranus Miller and Gurley, 1894, PI. 8, fig. 10, Raymond and
Barton, 1913, PI. 1, figs. 6-8, PI. 2, fig. 6, Slocom, 1913, PI. 17,
figs. 1-3, Caster, Dalve, and Pope, 1961, PI. 3, fig. 28 (only); C.
milleranus of Ludvigsen, 1979b, PI. 15, figs. 45-57, non Miller
and Gurley, 1894; C. parvilobatus Troedsson, 1928, PI. 17, fig.
14; C. pleurexanthemus Green, 1832, Evitt, 1953, PI. 5, figs. 1-
8, PI. 7, figs. 1,2, PI. 8, figs. 5-7; C. pleurexanthemus montyensis
Evitt, 1953, PI. 5, figs. 9-12, PI. 6, figs. 1-10, PI. 7, figs. 3, 5-
12, PI. 8, figs. 1-4, 8-16; C. ruidus Cooper, 1953, PI. 11, figs.
1,2,4, 12, 15, PI. 12, fig. 6, Shaw, 1974, PI. 7, figs. 1-10, 14,
15, and its synonyms (Shaw, 1974, p. 29); C. tuberosus Troeds-
son, 1928, PI. 18, figs. 1-9; and C. whittingtoni Evitt, 1953, PI.
5, figs. 13-16, PI. 6, figs. 11-24, PI. 7, figs. 4, 13-20, PI. 8, figs.
17-25, Whittington, 1992, PI. 2, figs. A-C, PI. 3, figs. A, B.
On the other hand, Gabriceraurus regroups Ceraurus blussoni
Ludvigsen, 1979b, PI. 13, figs. 9-29; C. dentatus Raymond and
Barton, 1913; C. cf. C. dentatus Raymond and Barton, 1913,
of Dean, 1979, PI. 3, figs. 1-7, PI. 4, figs. 1, 11; C. gabrielsi
Ludvigsen, 1979b, PI. 11, figs. 1-44, PI. 12, figs. 1-49, PI. 13,
figs. 1-8; C. hirsuitus Ludvigsen 1979b, PI. 14, figs. 1-25; and
C. plattinensis Foerste, 1920, PI. 21, figs. 20A, 20B, PI. 23, figs.
3A, 3B, DeMott, 1987, PI. 9, figs. 17-24, PI. 10, figs. 8-10,
Hessin, 1988b, fig. 1, A-D, Hessin, 1989, PI. 3, figs. 1-7. Ques-
tionably included are Gabriceraurus mijJlinensis DeMott, 1987,
PI. 9, figs. 8-16, PI. 10, figs. 4-7, and Ceraurus proicens Tripp,
1962, PI. 3, figs. 1-13.
Species of uncertain affinities. - Ceraurus maewestoides Lud-
vigsen, 1979b, PI. 13, figs. 30-33, C. binodosus Cooper and
Kindle, 1936, PI. 53, fig. 20, C. bituberculatus Troedsson, 1928,
PI. 17, fig. 12 (non fig. 11), and C. bispinosus Raymond and
Barton, 1913, PI. 1, figs. ?3, 4, Ludvigsen, 1978, fig. 26, all have
two swellings on their anterior glabellar lobes. Generic assign-
ment of these four species is uncertain, more so in view of the
fact that their respective pygidia are unknown. The pygidium
of Hapsiceraurus hispidus Whittington, 1954, PI. 59, figs. 1-7,
PI. 60, fig. 19, is equally unknown.
The specific assignment ofOntario specimens to Bufoceraurus
bispinosus (Raymond and Barton, 1913) by Hessin, 1989 (PI.
2, figs. 1-7) is probably in error.
Ceraurus tuberosus of Ludvigsen, 1979b, PI. 17, figs. 36-41,
and C. mackenziensis Ludvigsen, 1979b, PI. 15, figs. 1-44, can-
not be assigned unequivocally to either Ceraurus or Gabricerau-
rus, as they resemble Ceraurus in glabellar segmentation and
Gabriceraurus in the position of their eyes and the anterior
border.
SYSTEMATIC PALEONTOLOGY
Family REMOPLEURIDIDAE
Hawle and Corda, 1847
Subfamily REMOPLEURIDINAE
Hawle and Corda, 1847
Hypodicranotinae PRIBYL AND VANEK, 1972, p. 429.
Discussion. - As pointed out by Ludvigsen and Chatterton
(1991), the differences between the respective type genera of the
47-50, Ludvigsen, 1979b, PI. 13, fig. 9 xLI, 12 x 1.5,21 x 1.5,25 x 1.8; 51-55, Ludvigsen, 1979b, PI. 14, fig. 1 x 1.7,6 x 1.0,16 x 1.3,8
x 1.3, 12 x 1.2; 56-59, Raymond, 1921, PI. 11, fig. 8 x 0.6, PI. 10, fig. 3 x 0.4, herein, Fig. 5.3 x 0.6, Raymond, 1921, PI. 11, fig. 8 x 0.7; 60,
Raymond and Barton, 1913, PI. 1, fig. 2 xO.7; 61,63, DeMott, 1987, PI. 9, fig. 17 x 1.2, PI. 10, fig. 8 x 1.4; 62,64, Hessin, 1989, PI. 3, fig. 1
x 0.9, 4 x 1.5.
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7
Remopleuridinae and Hypodicranotinae are of degree and not
kind; Pribyl and Vanek's subfamily is thus best maintained as
a junior synonym of Remopleuridinae.
Genus HYPODICRANOTUS Whittington, 1952
Type species.-Remopleurides striatulus Walcott, 1875, by
original designation of Whittington, 1952, p. 1.
Discussion.-Ludvigsen and Chatterton's (1991) revision of
the genus has demonstrated the intraspecific variability of the
type species of the genus, as well as the wide geographic and
temporal (Llandeilian-Ashgillian) distribution ofthe genus. They
have synonymized the North American species Hypodicranotus
striatulus and H. missouriensis (Foerste, 1920); they recognized
only a single named species outside North America. The di-
agnosis of the genus must be modified in order to accommodate
evidence presented below as to the possession of 10 thoracic
segments, and not 11 as previously thought.
HYPODICRANOTUS STRIATULUS
(Walcott, 1875)
Figure 3.1-3.5
Remopleurides striatulus WALCOTI, 1875, p. 347, figs. 27A, 27a, 27b.
Hypodicranotus striatulus (Walcott). JOHNSON, 1985, unnumbered fig.
p. 94; LUDVIGSEN AND CHATIERTON, 1991, p. 620, figs. lA, IB, 2,
PI. 1, figs. 1-28 (complete synonymy).
Description. -As previously described by Whittington (1952)
and redescribed and modified by Ludvigsen and Chatterton,
1991, with the addition of the following.
Cranidium without ornamentation, occipital ring with barely
perceptible axial node. Maximum (exsag.) length to width of
hypostoma as 2.5: 1. Incomplete thorax with attached pygidium
retains six partial thoracic segments. Third before last segment
with stout sagittal spine, in posterior halfofaxial ring, projecting
posteriorly; width of spine at posterior margin of axial ring
approximately l/S width of axis at that level. Thorax smooth,
except for postero-Iaterally directed parts of thoracic pleurae
which are covered with terrace lines. Sagittal length ofpygidium
equal to maximum width occurring at mid-length; axis diffuse.
Surface of pygidium ornamented with terrace lines every 0.2
mm or so.
Discussion. -Johnson (1985, p. 94) had previously illustrated
Hypodicranotus striatulus with an axial spine on the eighth (from
the anterior) thoracic segment; although deformed, this Edenian
specimen from Ontario strongly suggests that the spine extends
to the pygidium. Ludvigsen and Chatterton's illustration (1991,
Fig. 1B) of a better preserved specimen from essentially the
same horizon unequivocally demonstrates the presence of a
stout axial spine on the eighth thoracic segment. As the pygidium
has but one segment, it is small, seldom preserved in individuals
retaining their thoraxes and, furthermore, mechanical prepa-
ration is consequently difficult; all this explains the uncertainties
as to the proper number of thoracic segments of this genus. The
Lac St-Jean specimen here illustrated (Figure 3.5) demonstrates
that the axial spine is on the third before last thoracic segment
and, consequently, the thorax of Hypodicranotus has 10 thoracic
segments, in contrast to the 11 in Remopleurides. Ludvigsen
and Chatterton's reconstructions (1991, fig. 2) ofthe type species
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 ORDOVICIAN TRILOBITES 9

of the genus with 11 thoracic segments thus has one thoracic
segment too many.
Material. - Four cranidia, 5 hypostomata, 2 librigenae, a par-
tial thoraco-pygidium and an isolated pygidium, from the Ship-
shaw (units 1 and 2, particularly abundant in the former) and
Des Galets Formations; also present in unnamed limestones at
Lake Manicouagan, Edenian. The most important localities are
1) quarries between Roberval and Pointe-Bleue, 1.5 and 2.2 km
north of the Ouiatchouaniche River in Roberval; 2) road out-
crops along route 169, east of Chambord; and, particularly, 3)
outcrops along the Shipshaw River, at the dam at Chutes aux
Galets and up to 1 km downstream, all UM collections.
Measurements. -All specimens with their exoskeletons, and
measurements are in mm. GSC 92974 hypostoma, sag. length
5.3, maximal exsag. length 18.7; GSC 11031 7 librigena, exsag.
length of eye: 3.6; GSC 110318 cranidium, sag. length: 4.7,
maximum width: 4.7; GSC 110319 pygidium, sag. length: 2.6,
maximum width: 2.4; and GSC 110320 incomplete thorax with
pygidium, sag. length of the 5 thoracic segments: 5.3.
Family ENCRINURIDAE Angelin, 1854
Subfamily ENCRINURINAE Angelin, 1854
Genus ENCRINUROIDES Reed, 1931
Type species. - Cybele sexcostata Salter, 1848, by original des-
ignation of Reed (1931, p. 20), as redescribed by Whittington,
1950.
Diagnosis. - Encrinurines with longer than broad glabella with
tips ofL2-L3-L410bes in straight line (exceptperiops); glabellar
tubercles small (tubercle size index 6-8), ranging from 40 (hor-
nei) to 200 (sexcostatus), with symmetrical tubercle pairs, except
in sexcostatus, across (tr.) Ll, L2 and L3; anterior cranidial
borders narrow (sag., exsag.), subhorizontal (hornei) to vertical
(sexcostatus), non-tuberculate (except periops); with short genal
spines and elongated middle hypostomal bodies (hypostomata
known in only two species); pygidia broader than long, axial
rings widely variable in number (8 in hornei, 12 in obesus, and
16 or 20 in the remaining two species), pleural ribs also variable
(5 in hornei, 6 inperiops, and 7 in remainder), and 1-2 congruent
pleural ribs and axial rings and 4 in hornei and obesus.
Species assigned. - Encrinuroides sexcostatus (Salter, 1848),
E. s.1. hornei (Dean, 1973), E. s.1. obesus Tripp, 1965, and E.
periops Tripp, 1967.
Discussion. -Even with the erection of new genera below, the
variability of Encrinuroides remains significant, as distinctive
suites of restricted characters define the other genera. This vari-
ability is probably due to its age span (Arenigian-Llanvirnian
to Ashgillian), the age limits of the taxa of Figure 1. This re-
stricted group containing Iapetus forms could possibly yield a
significant number of new species, in view of its duration, but
Iapetus related strata are restricted in occurrence.
Genus WALENCRINUROIDES n. gen.
Type species.-Ceraurus rarus Walcott, 1877, p. 68, Ray-
mond and Barton, 1913, p. 541, PI. 2, fig. 3, Chatterton and
Ludvigsen, 1976, p. 74, PI. 15, figs. 1-15, 16--43, non fig. 16,
DeMott, 1987, p. 80, PI. 11, figs. 13-25, non Tremblay and
Westrop, 1991, p. 821, fig. 17.27-17.36.
Diagnosis. - Encrinurines with variable width: length glabel-
lae and equally variable outlines of lines joining tips of L2-L3-
L4 lobes; glabellae with 50-90 tubercles, tubercle pairs sym-
metrical across (tr.) Ll and L2, and rarely L3 lobes, in some
species (lapworthi, rarus, and gelaisi n. sp.), without symmetry
in the others; short anterior cranidial borders in dorsal view,
with omnipresent tubercles; with short genal spines (except lap-
worthi) and hypostomata with elongated middle bodies (except
rarus); pygidia with subequal width-length ratios, 17-21 axial
rings, 7-9 pleural ribs, and 1-2.5 congruent pleural ribs and
axial rings.
Etymology. -Encrinuroides with a suffix of the first three let-
ters of the surname ofC. D. Walcott.
Species assigned.- Walencrinuroides rarus (Walcott, 1877);
W. s.l. gelaisi n. sp.; W. s.1. stincharensis (Reed, 1928); W. s.1.
lapworthi (Tripp, 1980b); W. autochthon (Tripp, 1962), W. po-
lypleura (Tripp, 1967), and W. n. sp. (=Encrinuroides rarus of
Tremblay and Westrop, 1991).
Discussion. - This is the only encrinurine of Figure 1 which
has subequal width: length pygidia, a character in itselfvariable
within certain species.
Walencrinuroides rarus has been described extensively and
illustrated by Chatterton and Ludvigsen (1976). Tremblay and
Westrop's (1991) specimens, referred to rarus, show a uniform
glabellar ornamentation, as pointed out by these two authors,
which is very distinct from the apomorphic concentration of
tubercles on the frontal lobe of other material referred to rarus
from the same general area as the type area of the species (see
DeMott, 1987).
Walencrinuroides gelaisi n. gen. n. sp. has not been chosen
as type of the genus because of its variability in pygidial shape
and the unknown nature of its hypostomal and rostral mor-
phology.
WALENCRINUROIDES s.l. GELAISI n. sp.
Figure 3.6-3.13
Encrinuroides n. sp. DESBIENS AND LESPERANCE, 1989, p. 1191.
Diagnosis. -A species characterized by reduced ornamenta-
tion, notably on the cephalic borders, thorax, and pygidial ribs.
Glabellar furrows short (tr.), distally restricted and sub-vertical;
cephalon with small genal spines or spinules. Pygidium as wide
as long to wider than long, with 19-20 tuberculated axial rings
and 7 (rarely, 8) p~eural ribs.

FIGURE 3-1-5, Hypodicranotus striatulus (Walcott, 1875).1, librigena, x 6.9, GSC 110317; 2, cranidium, x 6.0, GSC 110318; 3, pygidium, x 9.6,
GSC 110319; 4, hypostoma, x 3.0, GSC 92974; 5, incomplete thorax and pygidium with spine on third from posterior thoracic segment, x 6.7,
GSC 110320. 6-13, Walencrinuroides s.l. gelaisi new genus new species. 6-7, incomplete cranidium, dorsal and oblique lateral views, x 4.9,
x 4.4, GSC 110321; 8, incomplete cranidium, x 7.0, GSC 110322; 9, librigena, x 4.3, GSC 110323; 10-11, pygidia, x 8.1, x 4.6, GSC 110324,
110325; 12-13, holotype, enrolled individual, dorsal anterior and dorsal posterior views, x 5.9, GSC 110326. 14-19, Erratencrinurus s.l. spicatus
(Tripp, 1974). 14, incomplete cranidium, x 9.4, GSC 92988; 15, cranidium, with faint longitudinal glabellar furrow at arrow, glabellar tubercle
formula: Ll-l, L2-1,2 (on right doubled),3, F4-1,2,3, F4/5-0, F5-1,2, F6-1, x 7.0, GSC 110327; 16, librigena, x 6.4, GSC 110328; 17, pygidium,
x 7.0, GSC 92990; 18, cranidium, with an extra tubercle on the anterior border, left of sagittal line; glabellar tubercle formula: L 1-1, L2-1 ,2,3,
L3-1,2,3, F4-1 ,2,3, F-5, 1,2, x 6.1, GSC 110329; 19, incomplete cranidium with transverse L1-0 and larger L2-2 tubercles, x 9.4, GSC 110330.
Localities are: 4, Manicouagan Lake (see Desbiens and Lesperance, 1989); 5, 10, 50 m downstream from falls at Chute aux Galets, west side
of Shipshaw River, Chicoutimi outlier; remaining specimens from quarry on east side of road from Roberval to Pointe-Bleue, 1.5 km north
of Ouiatchouaniche River in Roberval, Lake St. John outlier.

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 10 LESPERANCE AND DESBIENS
Description. -Outline of cephalon hemispherical, maximal
cephalic width across genal spinules, twice cephalic length. Gla-
bella strongly convex sagittally, maximum height proximally
across L3; anterior glabellar lobe steeply sloping, remaining part
ofglabella sloping forward and backward evenly and much less
so. Maximal height of glabella, measured from axial furrows,
subequal to transverse fixigenal width proximally from eye lobe.
Maximal glabellar width across posterior part of anterior gla-
bella lobe, slightly wider than width across L3; width ofoccipital
ring slightly less than width across L3. Minimal glabellar width
across Ll, as 7: 10 when compared with maximalglabellarwidth.
Axial furrow incised and deep; preglabellar furrow frontally wid-
ened (sag., exsag.) at longitudinal median glabellar furrow, shal-
lower anterolaterally. Anterior border and field of fixigena sep-
arated by equally deep and wide furrow as preglabellar furrow,
originating at axial furrow-anterior border junction, joining lat-
eral border furrow of librigena; fossula not observed. Glabellar
furrows short (tr.), smooth, subvertical; S2 and S3 equally de-
veloped; SI more incised and shorter (exsag.) than S2-S3. Slope
lengths of glabellar furrows (tr.) and longitudinal median gla-
bellar furrow (sag.) essentially the same. Lateral glabellar lobes
equally developed transversely; L lone-half exsagittal length of
L2 and L3. Occipital furrow incised, transverse, on same line
(tr.) as proximal halfofposterior border furrow; posterior border
furrow incised, distal half anteriorly concave, dying out before
reaching posterior branch of facial suture. Both occipital and
posterior border furrows approximately equally long (sag., exsag.)
as deep. Occipital ring and posterior border almost smooth,
with rare 0.04-0.15 mm granules; posterior border shorter
(exsag.) than occipital ring distally. Eyes opposite S2 (tr.), pal-
pebral lobes sloping at 45°, at least as high as sagittal part of
Ll, smooth, delimited from field of fixigena by a sharp angu-
lation. Genal angles drawn out into spinules; posterior branch
of facial suture reaching margin immediately anteriorly of these
genal spinules.
Librigena with significantly elongated eye, of sloping length
half of total length of lateral border furrow and 3 times width
of same furrow (commonly measuring 1.6 by 0.9 mm); lateral
border furrow as wide as deep, bordered by numerous granules
proximally and distally, dying out posteriorly at faint sutural
ridge. Anterior border furrow shallow and wide, ill defined. Stalk
of eye lobe absent, junction of eye and field of librigena marked
by change ofslope, unfurrowed. Length twice width in both field
and precranidiallobe of librigena. Field of librigena tuberculate
and with numerous depressions. Lateral border little granulated;
anterior border of cephalon with a few tubercles or granules.
Rostral or median suture not preserved (GSC 110326 is laterally
and ventrally crushed).
Glabellar tubercles numerous, many (most ?) perforated, and
ca. 0.2 mm in diameter, some decidedly smaller and others
larger; tubercles Ll-l and L2-1 omnipresent; L2-2 also com-
mon, remaining tubercles showing no obvious distributional
pattern. Anterior border tubercles asymmetric in distribution,
commonly with 2 or 3 bigger tubercles on either half, with a
few smaller ones. Torular tubercle on fixigena surrounded by 8
or so slightly smaller tubercles arranged in circle; posteriormost
tubercle of circle is approximately exsagittal to torular one and
is the post-ocular tubercle. Depressions on field of fixigena and
librigena 0.1-0.2 mm in diameter. Fixigena and librigena with
0.2-0.4 mm tubercles. Tubercle size index 7.9-8.7.
Hypostoma unknown. Thorax with 11 segments. Axial rings
rounded, pleurae gently rounded, without exposed pleural fur-
rows. Axial rings and pleurae with 2 rounded inconspicuous 0.1
mm pustules.
Pygidium with subequal widths and lengths to wider than
long, in approximately equal proportions; greatest width slightly
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posterior of midpoint (sag.), outline lozengic with anterior sides
rounded. Pleural fields with 7 ribs, anteriormost 4 (? 5) ending
in free points, posteriormost one parallel to axis, and continuing
as a post-axial piece to slightly upturned posterior margin; bigger
individuals have better individualized seventh ribs with more
obvious and deeper axial furrows. Post-axial piece as long (sag.)
as posteriormost 5 axial rings. Two individuals with 8 pygidial
ribs; in both, seventh rib coalesces posteriorly sagittally and
continues to posterior margin, while what is here termed the
eighth rib forms post-axial ridge, equally long (sag.) as posterior-
most 5 axial rings and coalesced seventh rib posterior to eighth
rib, between posterior margin ofaxis and coalesced seventh ribs.
Pleural ribs rounded, unfurrowed, ranging from apparently un-
tuberculated and smooth to individuals with 1 or 2 low 0.1 mm
granules, distributed predominantly proximally. Axial rings 19-
20, last 4 or 5 indistinct to poorly individualized; central third
of axial ring furrows shallower than distal thirds, continuous in
only anteriormost 5 or 6; 2 congruent axial rings and pleural
ribs. Axial rings tuberculate, with 4 to 8 (or more) tubercles,
most prominent ones paired near sagittal line and 0.2 mm in
diameter, ranging in size downward to 0.05 mm, which tend to
be asymmetrically distributed.
Etymot%gy. -In honor of Philippe St-Gelais, of Jonquiere
in the Lake St. John district, a long-standing fossil collector in
this area.
Discussion. - The circle of eight or so bigger tubercles on the
fixigenae is reminiscent, but distinct from, the circlet of four
circumocular tubercles of Ramskold (1986; also, Edgecombe
and Chatterton, 1987), two of which are the torular and post-
ocular tubercles. Because of the presence of eight tubercles in
the Lake St. John species, these are not named circumocular.
The two individuals described above as possessing eight pygidial
pleural ribs are morphologically identical to Encrinuroides tho-
/us of Evitt and Tripp, 1977, PI. 4, fig. 4.
The other North American species of Wa/encrinuroides, W.
rarus and W. n. sp., have better developed glabellar segmen-
tation and fewer and coarser glabellar tubercles than does W.
s.l. ge/aisi n. sp. The highly tuberculated and rotund glabella of
this new species shows closest similarities with the North Amer-
ican species tho/us, capitonis, and the unnamed species of Ross
and Shaw, 1972. Wa/encrinuroides s.l. ge/aisi n. sp. is easily
distinguished from Ross and Shaw's (1972) species, which pos-
sesses a granulated-tuberculated posterior margin and forwardly
directed proximal glabellar and occipital furrows, which are
almost transverse in other North American species. The Okla-
homa species, capitonis, has cephalic borders more granulated
than in ge/aisi n. sp., it possesses more strongly developed gla-
bellar furrows, and its pygidium is segmented differently (14
axial rings and 8 pleural ribs). The closest affinities of ge/aisi n.
sp. are superficially, however, with tho/us. In addition to char-
acters detailed in Table 2, a notable difference lies in the amount
of granulation-tuberculation, which is far better developed in
tho/us than in ge/aisi n. sp., notably on the cephalic borders
and the whole of the pygidium. The glabellar tubercles of ge/aisi
n. sp. tend to be of the same size, but those in the central third
of the glabella of tho/us are bigger than the remainder.
Material. -One complete individual, 19 cranidia, 6 librige-
nae, and 27 pygidia, in various states of preservation, predom-
inantly with their exoskeletons, unit 1 of the Shipshaw For-
mation, Edenian, UM collections, from 1) quarry along the road
between Roberval and Pointe-Bleue, 1.5 km north of the Oui-
atchouaniche River in Roberval; 2) road outcrops along route
169, east of Chambord and 1.6 km west of Desbiens; 3) along
railroad, 350 m west of Chambord Junction; and 4) outcrops
along the Shipshaw River, immediately and up to 200 m down-
stream from the dam at Chute aux Galets.
 ORDOVICIAN TRILOBITES
Types. -An enrolled individual, GSC 110326, is herein des-
ignated holotype of the species; paratypes are GSC 110321-
110325.
Measurements. -All specimens with their exoskeletons, and
measurements are in mm. GSC 110321 incomplete cranidium,
sag. length ofglabella and occipital ring: 4.9, maximum glabellar
width: 3.9; GSC 110322 incomplete cranidium, as previous
measurements: 4.3 and 3.2; GSC 110323 librigena, maximal
length: 10.5; GSC 110324 incomplete pygidium, sag. length 4.4,
maximal width 6.4; GSC 110325 pygidium, as previous mea-
surements: 7.7, 8.6; GSC 110326 enrolled individual, cranidial
measurements as previously: 4.3 and 3.7, pygidium, sag. length,
including most of articulating half ring: 5.7, tr. width at, and
including, genal spines: 10.0.
Genus FRENCRINUROIDES n. gen.
Type species. - Encrinuroides capitonis Frederickson, 1964,
p. 71, PI. 1,figs. 1-5, Shaw, 1974,PI.9,figs.15, 19,20-23,PI.
10, figs. 1,2,4-11, ?non PI. 10, figs. 3, 12, 13.
Diagnosis. - Encrinurines with subequal width: length gla-
bellae and broader than long pygidia, variable tubercle size in-
dices (6-15), significant genal spines, round middle hypostomal
bodies with shallow rhynchos, 14-25 axial pygidial rings, 6-9
pleural ribs, and 1-3 congruent pleural ribs and axial rings.
Etymology. -Encrinuroides with a suffix ofthe first two letters
of the surname of E. A. Frederickson.
Species included. - Frencrinuroides capitonis (Frederickson,
1964), F. s.l. tholus (Evitt and Tripp, 1977), F. s.l. gibber (Dean,
1979), and F. torulatus (Evitt and Tripp, 1977).
Discussion. - Frencrinuroides n. gen. shares with Physema-
taspis longer genal spines than the other encrinurines of Figure
1, round middle hypostomal bodies, and pygidia that are broad-
er than long. Although it is not mentioned in the diagnosis, the
anterior cranidial borders are broadly intermediate in length
(sag., exsag., in dorsal view) between Physemataspis and Wal-
encrinuroides. On the other hand, Frencrinuroides shares with
Walencrinuroides adaxial tubercles on the fixigenae, and both
do not possess posterolateral hypostomal spines. Frencrinu-
roides is thus almost totally intermediate between these two
genera, as indicated by the cladogram of Figure 1.
Our knowledge of Frencrinuroides gibber is from Dean's type
material (1979, p. 10, PI. 5, figs. 8, 11, 12; PI. 6, figs. 2, 3) and
the associated cranidia that were identified as Ceraurus sp. (Dean,
1979, PI. 2, figs. 6, 7; PI. 4, figs. 3, 6, 9), which have been referred
to gibber by Edgecombe and Chatterton (1990, p. 823).
Evitt and Tripp (1977, p. 125) described their new species
Encrinuroides tholus as differing from E. torulatus in a number
of features. They did not give the pygidial width: length ratio
of tholus, and illustrated wider than long transitory pygidia and
two (holaspid) pygidia, one equally wide as long, and the other
wider than long. The pygidium of E. torulatus is decidedly wider
than long and thus available information suggests than the py-
gidium of tholus is wider than long (as coded in Table 2). It is
notable that if pygidial shape (character 17) of tholus is taken
as unknown in Table 2, exactly the same cladogram as Figure
1 is obtained, suggesting some type of robustness in the data.
The type species of Frencrinuroides n. gen. is taken to be
capitonis, in preference to tholus, as the latter is rare as an adult
(Evitt and Tripp, 1977, table 1), and its adult genal angle, li-
brigena, hypostoma, and thorax are unknown.
Genus PHYSEMATASPIS Evitt and Tripp, 1977
Type species. - Physemataspis coopi, Evitt and Tripp, 1977,
p. 138, PI. 12, figs. 1-4, PI. 13, figs. 1-17, PI. 14, figs. 1-7, text-
figure 12, 13, by original designation.
Diagnosis. - Encrinurines with variable glabellar outlines, high
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11
tubercle size indices (10-15), significant genal spines, long (exsag.,
generally also sag.) cranidial anterior border in dorsal view,
commonly tuberculate, or at least coarsely granulose, with round
middle hypostomal bodies (except in neuter) and posterolateral
hypostomal spines (except in insularis), and broader than long
pygidia with 13-18 axial rings, 6-8 pleural ribs, and 2-3.5 con-
gruent pleural ribs and axial rings.
Species assigned. -Physemataspis (Physemataspis) coopi Ev-
itt and Tripp, 1977, P. (Physemataspis) mirabilis Tripp, 1980a,
P. (Prophysemataspis) n. subgen. uncatus (Evitt and Tripp, 1977),
P. (Prophysemataspis) n. subgen. neuter (Evitt and Tripp, 1977),
and P. s.1. insularis (Shaw, 1968).
Discussion. - The semicircular glabellae ofthree ofthe species
included within this genus are unique among the encrinurines
treated here, as are the hypostomata with posterolateral spines
and round middle bodies, the absence of adaxial tubercles on
the fixigenae, and the long (exsag., commonly also sag., in dorsal
view) anterior cranidial borders, approximately equal or slightly
less than the sag. lengths of the occipital rings.
As the three species with semicircular glabellae in this genus
are visually distinctive, so are the species neuter and uncatus,
shown through phylogenetic analysis to belong to the Physe-
mataspis clade, when compared to these three species. This
morphological distinctness, reinforced by three character changes
at the neuter-uncatus node (Figure 1), are taken as sufficient
justification to separate these two species taxonomically at the
subgeneric level, as P. (Prophysemataspis) n. subgen.
A sagittal depression on the frontal lobe of the glabella is
present in Encrinuroides (except E. periops), Walencrinuroides
n. gen., Frencrinuroides n. gen., and Physemataspis (Prophyse-
mataspis) n. subgen., but is absent in P. (Physemataspis).
Subgenus PHYSEMATASPIS (PHYSEMATASPIS)
Type species. -As for the genus.
Diagnosis. -Glabellae subcircular, without a sagittal depres-
sion on the frontal lobe; pygidia with 13 axial rings, 6-7 pleural
ribs, and paired sagittal axial tubercles.
Discussion. - The anterior cranidial border is absent sagittally
in P. (Physemataspis) mirabilis and reduced in P. (Physema-
taspis) coopi; this last species is also unique within the two
subgenera in possessing axial thoracic spines (however, the state
of this character is unknown in the other species). The hypos-
toma is only known in P. (Physemataspis) coopi and it is without
rhynchos.
Subgenus PHYSEMATASPIS (PROPHYSEMATASPIS)
n. subgen.
Type species. - Encrinuroides uncatus Evitt and Tripp, 1977,
p. 128, figs. 4A, 8, PI. 5-7.
Diagnosis. -Glabellae with straight or concave outwards out-
lines, 13-15 tubercle size index, and L 1-L2-L3 glabellar tuber-
cle pairs; hypostomata with rhynchos; pygidia with 14-18 axial
rings, 7-8 pleural ribs, and sagittal axial tubercles.
Etymology. - From Physemataspis and the latin suffix pro,
generally translated as meaning/or.
Discussion. - The symmetry and coarse ornamentation on the
glabella and the cranidial anterior border within this subgenus
suggest affinities with Erratencrinurus, a subject best investi-
gated in a separate phylogenetic analysis.
Genus ERRATENCRINURUS Krueger, 1971
Type species. - Erratencrinurus capricornu Krueger, 1971, p.
1147, PI. 6, figs. 1, 2, PI. 7, figs. 1, 2, by subsequent designation
of Krueger, 1972, p. 858.
 12 LESPERANCE AND DESBIENS
ERRATENCRINURUS s.l. SPICATUS (Tripp, 1974)
Figure 3.14-3.19
Encrinurus spicatus TRIPp, 1974, p. 485-487, PI. 1, figs. 1-8.
Encrinuroides? spicatus (Tripp). DESBIENS AND LESPERANCE, 1989, figs.
2Q-2S.
Erratencrinurus s.1. spicatus (Tripp). OWEN AND HEATH, 1990, p. 228.
Description. -As described by Tripp (1974), except for the
following. Anterior cranidial border always bisected by median
furrow, with eight tubercles; one specimen (Fig. 3.18) has ninth
tubercle that is, however, not in sagittal plane, and thus asym-
metric. Longitudinal median glabellar furrow ranging from ab-
sent to incised, of sag. length subequal to exsag. length between
glabellar lobes; relative proportion between presence or absence
of median glabellar furrow unknown. Lateral (glabellar) lobes
pointed tubercles; interspaces between these lobes without (gla-
bellar) furrows as such, but concave. Four specimens have larger
glabellar tubercles, as previously described by Tripp (1974), but
in one specimen (Fig. 3.14) Ll-l and L2-2 are larger and only
L2-2 is larger in another specimen (Fig. 3.19), which also has a
transverse L 1-0 tubercle.
Pygidium with well-developed articulating half ring and half
ribs. Sixth and posteriormost pleural rib parallel to axis; height
of rib significantly less than sagittal height of axis.
Discussion. - Differences between the Lake St. John material
and that originally described from Wisconsin can be easily ex-
plained by differences between two small samples and normal
geographic variation present in any species. As such, these dif-
ferences do not warrant subspecific (and much less specific)
separation. Even within the small sample here studied, the vari-
ation in glabellar tubercles is best interpreted as inherent to the
species. It has, however, been suggested that such variation is
the norm in some Silurian encrinurines (Ramskold, 1986, p.
529).
A pygidium with eight pleural ribs, otherwise identical to
spicatus, is not assigned to this species. This pygidium from a
locality on the Shipshaw River does not occur with other en-
crinurine material, and this renders its specific assignment prob-
lematical.
The presence of a median glabellar furrow in this species had
not been noted previously. Its occurrence links spicatus with the
species of Figure 1, with the exception of Physemataspis (Phy-
semataspis), reinforcing previous suggestions ofits intermediate
phylogenetic position.
Material. - Eight cranidia, 7 pygidia and 14 librigenae, all
with preserved exoskeletons and from unit 1 of the Shipshaw
Formation (Edenian-Upper Ordovician), UM collections, from
1) quarry along the road between Roberval and Pointe-Bleue,
1.5 km north ofthe Ouiatchouaniche River in Roberval; 2) road
outcrops along route 169, east of Chambord and 1.6 km west
of Desbiens; 3) along railroad, 350 m west ofChambord Junc-
tion; and 4) outcrops along the Shipshaw River, immediately
and up to 1 km downstream from the dam at Chute aux Galets.
Also present in the Manicouagan Lake outlier area (GSC col-
lections).
Measurements. -All specimens with their exoskeletons, and
measurements in mm. GSC 110327, 92988, 110329, and 110330
incomplete cranidia, sag. length of glabella and occipital ring,
respectively, 3.1, 3.3, 3.6, and 3.1, maximal glabellar width,
respectively, 2.7, 2.5, 3.0, and 2.5, and tr. width of occipital
ring ofGSC 110329: 2.6; GSC 110328librigena, maximal length:
6.9; and GSC 92990 pygidium, sag. length: 4.5, maximal width:
5.1.
Family CHEIRURIDAE Hawle and Corda, 1847
Remark. - The justification ofthe familial authorship is given
by Lane, 1971, p. 7.
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Subfamily CHEIRURINAE Hawle and Corda, 1847
Genus CERAURUS Green, 1832
Ceraurus (Eoceraurus) ESKER, 1964, p. 201.
?Hapsiceraurus WHITTINGTON, 1954, p. 125.
?Bufoceraurus HESSIN, 1989, p. 1208.
Type species. - Ceraurus pleurexanthemus Green, 1832, p. 83,
by monotypy.
Diagnosis. -A cheirurine genus with forwardly expandinggla-
bella widest across L3 and/or posterior part of anterior lobe,
eyes transverse to S2 and/or L3, with steeply sloping to vertical
anterior borders, and preglabellar furrow merging into axial
furrow; glabellar segmentation reduced so that central two-thirds
(or more) of glabella, tr. of L2 and L3, unsegmented; glabellar
ornamentation ranging from symmetrical to disorganized, with
tendency to symmetrical; pygidium essentially twice as wide as
long; pygidial spines uncarinated; segmentation proximally of
great spines with subtriangular posterior margin.
Discussion. - In dorsal view, the glabella reaches, and even
overhangs, the anterior margin, as the anterior border is vertical.
Natural populations of this genus can have eyes that are both
transverse to L3 and S2, and there is a tendency for the indi-
viduals to have eyes transverse to L3; on the other hand, Cerau-
rus milleranus appears to have eyes exclusively opposite S2.
Whittakerites Ludvigsen, 1976, has eyes transversely from S1
and a truncated semi-elliptical glabella, which serve to distin-
guish it from Ceraurus.
The carinated great pygidial spines and subrectangular and
smooth glabella of Ceraurinus Barton, 1913 (Ludvigsen, 1977),
clearly differentiate this genus from Ceraurus. Ceraurinella Coo-
per, 1953, is somewhat similar, but its pygidial spines are not
carinated; its glabella is also subrectangular and in this case
granulose. The bulb-shaped glabella of Borealaspis Ludvigsen,
1976, widest across L3, and possessing a posterior glabellar (sag.)
spine, is distinctive when compared to Ceraurus; its palpebral
lobes across L3 or S3 recall Ceraurus and its spinose pygidium
is almost identical to some species of Gabriceraurus, testifying
to the close affinities of all these cheirurine taxa.
Hapsiceraurus hispidus Whittington, 1954 (the species being
the type of the genus, both erected in the same publication),
from the Canadian Arctic, is known only from individuals with-
out pygidia. Lane (1971, p. 75) referred to a pygidium (of un-
known origin) which possibly belongs to the genus. Until the
pygidium of Hapsiceraurus is definitely identified, the genus is
best considered a possible synonym of Ceraurus, from which it
differs in few respects.
Both Lane (1971) and Shaw (1974) did not consider Ceraurus
(Eoceraurus), with C. (Eoceraurus) trapezoidalis Esker, 1964,
as type species, a valid subgenus; furthermore, trapezoidalis
itself has been synonymized with C. ruidus Cooper, 1953, by
Shaw (1974, p. 29).
Bufoceraurus of Hessin, 1989, has for type species the very
poorly known Ceraurus bispinosus Raymond and Barton, 1913,
based on an incomplete holotype cranidium. Hessin's (1989)
material does not show the distinctive spines, or protuberances,
on the frontal glabellar lobe, and his assignment of Ontario
material to this species is questionable. In such circumstances,
the taxon bispinosus is best restricted to the holotype, and Bufo-
ceraurus considered a possible subjective synonym of Ceraurus.
Species included. - Ceraurus cetus Dean, 1979; C. globulo-
batus Bradley, 1930; C. cf. C. globulobatus of Hessin, 1989; C.
matranseris Sinclair, 1947; C. milleranus Miller and Gurley,
1894; C. milleranus of Ludvigsen, 1979b; C. parvilobatus
Troedsson, 1928; C. pleurexanthemus Green, 1832; C. pleurex-
anthemus montyensis Evitt, 1953; C. ruidus Cooper, 1953; C.
tuberosus Troedsson, 1928; and C. whittingtoni Evitt, 1953.
 ORDOVICIAN TRILOBITES
CERAURUS GLOBULOBATUS Bradley, 1930
Figure 4.14-4.23
Ceraurus globulobatus BRADLEY, 1930, p. 274, PI. 30, figs. 33-36, 39-
42, ?fig. 38.
non Ceraurus cf. globulobatus Bradley. HESSIN, 1989, p. 1217, PI. 4,
figs. 1-7.
Diagnosis. - A species of Ceraurus with bituberculate (tuber-
cles of 0.3 and 0.6 mm) glabella with commonly an undefined
pattern of tubercle distribution to one in which two ill-defined
exsagittal rows of 5-6 coarser tubercles can be recognized, an
occipital node, and glabellar furrows equally incised, essentially
transverse; largest specimens have fewer smaller glabellar tu-
bercles; maximum width of glabella across posterior part of
frontal lobe, only slightly greater than across L3; frontal glabellar
lobe laterally steeply sloping, and even overhanging axial furrow
on inner molds; center (exsag.) of palpebral lobe opposite (tr.)
L3 and S2; lateral border furrow turning transversely at anterior
end, joining S3; eye ridge absent to ill developed, low, and
forming part of posterior slope of proximal part of transverse
prolongation of lateral border near S3; distal part of eye ridge
near eye unknown (or possibly not developed); fixigenae retic-
ulate and with small tubercles, notably near posterior border
furrow, and two large tubercles disposed exsagittally between
eye and glabella aligning themselves with a third tubercle on
posterior border; anterior border visible dorsally, one-third to
one-half as long (sag.) as occipital ring, smooth or bituberculate
with scarce tubercles. Hypostoma maculate, equally wide as
long, with a steeply sloping (not vertical) anterior border and a
slightly dorsally deflected posterior border. Pygidium with small
projections between the great spines, posterior margin trian-
gular; axial furrows between axis and pleural ribs or spines
obsolete.
Types. - Holotype, inner mold of incomplete cranidium, DC
20709A; paratype, inner mold of large incomplete cranidium,
DC 20719A; paratype, incomplete pygidium retaining part of
exoskeleton, DC 20695C; paratype, incomplete outer mold of
pygidium, UC 20695B; paratype, inner mold ofan almost com-
plete hypostoma, DC 20709B; and hypotype, incomplete inner
mold of cranidium, UC 28956 (fig. 33 of Bradley, 1930).
Material. -All the material of Ceraurus globulobatus exam-
ined from the Field Museum (UC) is from the middle Ordo-
vician (Kirkfieldian-Shermanian: Ross et aI., 1982) Kimmswick
Limestone of Missouri (Glen Park-Sulphur Springs area; rarely,
Festus) and Illinois (1 mile north of Batchtown). This material
is composed almost exclusively of inner molds and totals ap-
proximately 55 cranidia, 7 fixigenae, 6 pygidia, and 33 hypos-
tomata.
Discussion. - The extended diagnosis above, in conjunction
with Bradley's (1930) description, is sufficient to characterize
the species. It should be noted that the concept of the species
rests almost exclusively on inner molds, with consequent un-
certain differences on the external surface.
Paratype UC 20695A (original of PI. 30, fig. 38 of Bradley,
1930), inner mold ofincomplete cranidium, with missing frontal
part of glabella, as well as another cranidium from UC 5869
(Glen Park, Missouri) are assigned doubtfully to Ceraurus glob-
ulobatus because of their flattened profiles (sag., tr.), contrasting
with the evenly rounded profiles of C. globulobatus.
Ceraurus globulobatus differs from the type of the genus, C.
pleurexanthemus, by the lateral slope of the frontal lobe, which
is far more gentle (45°, or so) in the latter (but this feature has
not been determined with precision for all species of the genus).
Ceraurus globulobatus differs from milleranus of Ludvigsen,
1979b, pleurexanthemus, pleurexanthemus montyensis, ruidus,
tuberosus, and whittingtoni by its dorsally visible frontal border.
Ceraurus cf. C. globulobatus of Hessin, 1989, has a dorsally
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13
visible frontal border, its maximal glabellar width across the
posterior part of the frontal glabellar lobe is distinctly greater,
and it has more strongly developed eye ridges than does glob-
ulobatus (in fact, stronger eye ridges are common to almost all
species of Ceraurus). The hypostoma of globulobatus apparently
differs from all other hypostomata referred to Ceraurus by its
equal width and length, and its anterior and posterior borders,
which have different dorso-ventral inclinations. The two tu-
bercles on the fixigenae, aligned with a third one on the posterior
border, are common in Ceraurus, but they do not occur in
ruidus; much the same is true for the common occurrence of
two exsagittal rows of tubercles on the glabella, but these rows
are absent in ruidus, tuberosus, and whittingtoni. Ceraurus cetus
is unique in possessing two spines proximally ofthe genal spines.
Differences between globulobatus and matranseris are detailed
below.
Measurements. -All specimens are inner molds, except UC
20685C an outer mold, and measurements are in mm. UC
20709a, 29103, 28956, 20719b, and 20696 incomplete cranidia,
sag. length of glabella and occipital ring, respectively, 10.7, 6.7,
9.3, 16.1, and> 5.3, maximal glabellar width, respectively, 8.5,
5.6,7.2, 12.8, and 4.6. UC 20709 hypostoma, sag. length: 5.8.
UC 20695c, 20695b, pygidia, sag. length, respectively, 4.9 and
3.7.
CERAURUS MATRANSERIS Sinclair, 1947
Figure 4.1-4.13
Ceraurus matranseris SINCLAIR, 1947, p. 254, PI. 1, figs. 3-6; DESBIENS
AND LESPERANCE, 1989, p. 1191, fig. 2M, 2N.
Diagnosis. - A species of Ceraurus with bituberculate (tuber-
cles of0.3 and 0.6-1.0 mm) glabella with almost always a pattern
of tubercle distribution of two exsagittal rows of 5-6 coarser
tubercles, occipital node minute, present only in a few individ-
uals, and essentially transverse S2 and S3 glabellar furrows, S2
shallower; some individuals without smaller glabellar tubercles
and others with tubercle on each glabellar lobe; maximum width
of glabella across posterior part of frontal lobe; frontal glabellar
lobe laterally sloping at 45°-60°;center (exsag.) ofpalpebral lobes
commonly opposite (tr.) L3, rarely S2; lateral border furrow
turning transversely and slightly posteriorly at its anterior end,
joining S3; eye ridges immediately posterior of transverse part
of lateral border furrow, essentially transverse to S3, a few in-
dividuals have eye ridges slightly anteriorly directed proximally;
fixigenae reticulate and with few small or large tubercles, and
two large tubercles disposed exsagittally between eye and gla-
bella, in line with third tubercle on posterior border; anterior
border not visible to visible dorsally, pustulose and bituber-
culate with scarce tubercles. Maculae on hypostoma faintly pres-
ent in a few individuals; hypostoma much wider than long (as
3:2), without anterior border in central halfand slightly dorsally
upturned posterior border. Missing hypostomal anterior border
slightly more than half as wide as anterior lobe of hypostoma.
Rostral plate as wide as anterior lobe of hypostoma. Cephalic
doublure pustulose. Thorax with two tubercles on each axial
ring. Pygidium with triangular posterior margin; axial furrows
between axis and pleural ribs or spines obsolete to faint.
Discussion. - Ceraurus matranseris and C. globulobatus have
been considered possible synonyms (Desbiens and Lesperance,
1989, table 1). The diagnoses of these two species show that in
fact they are quite distinct, even though globulobatus is essen-
tially known from inner molds and the exoskeleton of matran-
seris is well known. Nonetheless, they are close to one another;
matranseris is, however, younger (Edenian). Both species show
significant variation, as detailed. The better-developed eye ridg-
es, the shallower lateral slope of the frontal glabellar lobe and
the symmetrical tubercles near the sagittal line of the glabella
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 ORDOVICIAN TRILOBITES 15

clearly distinguish C. matranseris from C. globulobatus. These
symmetrical glabellar tubercles are more obvious (in size and
symmetry) in matranseris than in other species of Ceraurus (for
further discussion, see the previous discussion of C. globulo-
batus).
The hypostoma of Ceraurus matranseris is very similar to the
hypostomata of C. pleurexanthemus, C. pleurexanthemus mon-
tyensis, and C. whittingtoni of Evitt (1953, PI. 7, figs. 1-20), but
the Quebec material is not silicified and it is impossible to
confirm that it does (or does not) possess the short frontal margin
at right angle to the surface of the hypostoma. Perhaps this
feature is variable, as C. whittingtoni illustrated by Whittington
(1992, PI. 2, fig. B) has an anterior hypostomal margin sloping
anteriorly. The dorsally upturned hypostomal posterior margin
of C. matranseris (also present in C. globulobatus) is absent
from Evitt's (1953) material. Maculae are well developed in C.
pleurexanthemus and C. pleurexanthemus montyensis, but faint
or absent in C. whittingtoni and C. matranseris. The hypostoma
of C. ruidus is distinctive as it is more subrectangular, with
better developed anterior lateral borders, than the hypostomata
discussed above.
Material. - This species is known from specimens with their
exoskeletons, which include, approximately, 2 complete indi-
viduals, 1 cephalon, 50 cranidia, 6 glabellae, 6 librigenae, 13
hypostomata, and 7 pygidia, predominantly from unit 1 of the
Shipshaw Formation, but also from unit 2 ofthe same formation
and the Des Galets Formation (UM collections) and the Man-
icouagan Lake area (GSC collections), all of which are Edenian.
Localities from the Lake St. John district include 1) quarries
along the road between Roberval and Pointe-Bleue, 1.5 and 2.2
km north of the Ouiatchouaniche River in Roberval; 2) road
outcrops along route 169, east of Chambord and 1.6 km west
of Desbiens; 3) along railroad, 350 m west ofChambord Junc-
tion; 4) outcrops along the Shipshaw River, immediately and
up to 1 km downstream from the dam at Chute aux Galets; and
5) drainage ditch, 200 m east of the Niobec road in St-Honore.
The quarry 1.5 km north of Roberval has yielded abundant
material, as has another locality in unit 1 in the Shipshaw River
some 250 m downstream from the dam.
Types. - Holotype, small cranidium GSC 6801, paratype, in-
complete pygidium GSC 7395, pathological individual, large
cranidium GSC 7712 all of Sinclair, 1947, from quarries "1
mile north of Roberval" (p. 255), and hypotypes GSC 92984,
92985 of Desbiens and Lesperance, 1989.
Measurements. -All specimens have their exoskeletons, and
measurements are in mm. Glabellar and occipital ring sag. lengths
of GSC 92985, 110332, 110333, 6801, and 92984 are, respec-
tively, 7.4, 9.3,8.2,2.9, and 9.1, while the maximum glabellar
widths are, respectively, 6.4, 7.4, 7.4, 2.6, and 8.5. GSC 92985
and 110333 have tr. widths across the palpebral lobes of 14.9
and 16.2. The articulated specimen GSC 92984 has, further-
more, a dorsal sag. length of 33.4, and a hypostomal sag. length
of 6.7. Pygidia GSC 7395 and 110334 have, respectively, sag.
lengths of2.0 and 4.2, and tr. widths anteriorly are 5.2 and 10.0.
Genus GABRICERAURUS Pribyl and Vanek, 1985
Type species. - Ceraurus gabrielsi Ludvigsen, 1979b, by orig-
inal designation of Pribyl and Vanek, 1985, p. 161.
Diagnosis. -A cheirurine genus with subrectangular to for-
wardly expanding glabella widest across posterior part of an-
terior glabellar lobe, eyes transverse from S 1 to S2, more com-
monly L2, with distinct anterior border in dorsal view, and
incised preglabellar furrow merging with axial furrows; glabellar
segmentation significant, central halfacross (tr.) L2 and L3 with-
out segmentation; glabellar ornamentation with strong tendency
to asymmetricity; pygidial segmentation proximally of great
spines with tendency to spinose transverse margins, approxi-
mately twice as wide as long.
Species included. - Ceraurus blussoni Ludvigsen, 1979b; C.
dentatus Raymond and Barton, 1913; C. cf. C. dentatus of Dean,
1979; C. gabrielsi Ludvigsen, 1979b; C. hirsuitus Ludvigsen,
1979b; C. plattinensis Foerste, 1920; ?Gabriceraurus mifJlinensis
DeMott, 1987; and ?Ceraurus proicens Tripp, 1962.
Discussion. -See Ceraurus above.
GABRICERAURUS DENTATUS (Raymond and Barton, 1913)
Figure 5.1-5.3
Ceraurus pleurexanthemus Green, 1832. HALL, 1847, p. 242, PI. 65, fig.
Id-lf, Ih, Ii, lk, ?11, 1m, ?In; PI. 66, fig. la-lh; non PI. 65, fig. la-
Ic, Ig.
Cheirurus pleurexanthemus (Green). BILLINGS in I...cxJAN, MURRAY, HUNT,
AND BILLINGS, 1863, p. 188, fig. 188.
Ceraurus dentatus RAYMOND AND BARTON, 1913, p. 534, PI. 1, fig. 2,
PI. 2, figs. 4, 5; RAYMOND, 1921, PI. 10, figs. 2, 3, PI. 11, figs. 7, 8;
DEMolT, 1987, PI. 9, figs. 1-7, PI. 10, figs. 1-3.
non Ceraurus dentatus Raymond and Barton. RAYMOND, 1921, PI. 10,
fig. 2; SINCLAIR, 1964, PI. 4, fig. 10; LUDVIGSEN, 1978, fig. 33; LUDV-
IGSEN, 1979a, fig. 26C.
Ceraurus hirsuitus Ludvigsen. HESSIN, 1989, PI. 1, figs. 1-7.
Primary types assigned.-Paratypes AMNH 29511, MCZ 7,
GSC 1769b, and 8062.
Primary type rejected. -Paratype GSC 1769 (=non Ceraurus
dentatus of synonymy list).
Other material. - Extensive material from the "Trenton" at
Middleville, New York, illustrated by Hall (1847) was previ-
ously catalogued in the American Museum of Natural History
under the all inclusive (locality) number 850/1 and has been
renumbered. A single specimen from this lot is a paratype (see
below; AMNH 29511), while the remainder are hypotypes of
dentatus. These are: 29502 (PI. 65, fig. Id, Ie); 29503 (PI. 65,
fig. If); 29505, 29506 (PI. 65, fig. Ih, Ii, respectively); 29507-
29510 (PI. lk-ln, respectively); 29512 (PI. 66, unnumbered
figure between multiple views of 29511).
Description. -As in DeMott, 1987, p. 78, with addition of the

FIGURE 4 -1-13, Ceraurus matranseris Sinclair, 1947. 1-3, cranidium, oblique anterolateral, lateral, and dorsal views, x 3.5, x 3.5, x 2.3, GSC
92985; 4-5, incomplete individual, oblique anterolateral and dorsal views, x 3.1, x 2.7, GSC 110331; 6-7, incomplete cranidium, dorsal and
frontal views, x 3.0, x 4.7, GSC 110332; 8, cranidium, x 2.3, GSC 110333; 9, ho1otype cranidium, x 6.7, GSC 6801; 10-11, almost complete
individual, dorsal and ventral cephalic views, posterior pygidial spines exagerated by mechanical preparation, x 2.2, x 2.8, GSC 92984; 12,
paratype pygidium x 4.5, GSC 7395; 13, pygidium, x 2.7, GSC 110334.14-23, Ceraurus globulobatus Bradley, 1930.14-15, holotype cranidium,
dorsal and lateral views, x 2.4, x 4.0, UC 20709A; 16, cranidium, x 3.4, UC 29103; 17, Bradley, 1930, hypotype cranidium, x 2.7, UC 28956;
18, paratype hypostoma, x 3.8, UC 20709B; 19, paratype incomplete cranidium, x 1.6, UC 20719A; 20-21, incomplete cranidium, dorsal and
oblique anterolateral view, x4.6, x 8.1, UC 20696; 22, paratype pygidium, x 3.9, UC 20695C; 23, paratype pygidium, x4.9, UC 20695B.
Localities are: 1-3, Manicouagan Lake; 4, 5, 10, 11: same quarry detailed in explanation of Figure 3; 6-8,13: Shipshaw River as detailed in
explanation of Figure 3; 9, 12: see Sinclair, 1947; 14-18, Glen Park-Sulphur Springs area, Illinois; 19-23: 1 mile north of Batchtown, Illinois.

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 16 LESPERANCE AND DESBIENS

FIGURE 5- Gabriceraurus dentatus (Raymond and Barton, 1913). 1, paratype, incomplete individual with exoskeleton, x 1.0, Middleville, Herkimer
County, New York, AMNH 29511, original of Hall, 1847, PI. 66, fig. la-Ih; 2, hypotype, incomplete pygidium with exoskeleton, x 1.0, same
locality as 1, AMNH 29509, original of Hall, 1847, PI. 65, fig. 1m; 3, paratype, almost complete individual with exoskeleton, Cobourg, Ontario,
x 1.0, GSC I769b, original of Raymond, 1921, PI. 10, fig. I.

following. Entire surface of exoskeleton covered with 0.08--0.1
mm granules. Genal spines exceptionally long (exsag.) for the
genus, covering two-thirds of thoracic length.
Discussion.-Hessin (1989, p. 1207) has used the taxon hir-
suitus Ludvigsen, 1979b, as synonymous with dentatus Ray-
mond and Barton, 1913, and as a replacement name for the lost
holotype of dentatus (Bolton, 1966, p. 50). Furthermore, he
stated that there is a single existing paratype pygidium (MCZ
7), and that it was not designated a neotype in view of the
difficulty in using solely pygidia in (cheirurine) taxonomy. This
unusual procedure ofa "replacement" name has no justification
in the Code of Zoological Nomenclature (Ride et aI., 1985).
What is perhaps more serious is that additional paratypes are
extant besides MCZ 7 and a neotype is unwarranted and un-
justified.
Raymond and Barton's (1913) description oftheir new species
dentatus (repeated in Raymond, 1921) includes a synonymy, a
short description, and a discussion of other specimens besides
the explicit erection of a holotype (GSC 1775, a complete in-
dividual, now lost). According to the Code, all the specimens
mentioned become part of the type series (Article 72(b)(i» and
as a holotype was erected the other specimens become paratypes
(and not syntypes, and thus unavailable for lectotype selection)
(Article 72(b)(v». The concept of the taxon dentatus must con-
sequently rest on any or all of the paratypes and a replacement
name is illegal and unjustified. Although dentatus and hirsuitus
Ludvigsen, 1979b, are very similar, they are not considered
synonymous, as hirsuitus has far longer (exsag.) palpebral lobes
(and hence, eyes) and a different course of the anterior branch
of the facial suture, with a consequent greater fixigena proxi-
mally and anteriorly of the eye in dentatus.
In order to stabilize the concept of dentatus, the extant para-
types must first be ascertained. The following paratypes have
been traced, or examined: AMNH 29511 (illustrated by Hall,
1847, PI. 66, fig. la-lh; herein, Figure 5.1), an incomplete in-
dividual with missing posterior part of thorax and pygidium,
MCZ 7 (illustrated by Raymond and Barton, 1913, PI. I, fig.
2), a pygidium, and GSC 1769, 1769b (herein, Figure 5.3), and
8062, all illustrated on PI. 10 of Raymond 1921, which can be
identified from Raymond and Barton's (1913) discussion (more
or less complete individuals).
GSC 1769 is herein excluded from dentatus as it possesses
eyes that are set far more forward (tr. of L2 or S2) than in the
remaining paratypes; furthermore, its glabella recalls Ceraurus
pleurexanthemus.
GSC 1769b is well preserved, but fragile. It is herein reillus-
trated and it is suggested that the concept of dentatus be derived
predominantly from this specimen, as it appears closest to the
lost holotype. MCZ 7 is a pygidium that appears identical to
the holotype, and it need not be reillustrated. Figure 5.2 is a
hypotype pygidium from the same locality as paratype AMNH
29511.
ACKNOWLEDGMENTS
The writers are grateful to the following individuals and their
respective institutions for the loan and information ofboth type
and non-type material: T. E. Bolton and J. Dougherty of the
Geological Survey of Canada, Ottawa (GSC), F. Collier of the
Museum of Comparative Zoology, Harvard University (MCZ),
N. Eldredge and C. B. Boyko of the American Museum of Nat-
ural History (AMNH), and C. Foster and S. Lidgard ofthe Field
Museum of Natural History (presently UC, WM of Bradley,

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 ORDOVICIAN TRILOBITES
1930). M. Bizzarro, G. D. Edgecombe, F. C. Shaw, and S. Wes-
trop provided helpful comments on the contents of this contri-
bution. Past and present grants of the National Sciences and
Engineering Research Council of Canada to PIL have proven
essential and are appreciated. UM collections refer to the non-
type collections of the Departement de geologie, Universite de
Montreal, Montreal.
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ACCEPTED 1 MAY 1995
The Faculte des Arts et des Sciences of the Universite de Montreal
has paid for this contribution.
 ORDOVICIAN TRILOBITES
APPENDIX
The following characters were used in the phylogenetic analysis.
1) Sagittal depression on anterior part of glabella; 0 = presence; 1 =
absence.
This falls within Temple and Tripp's (1979) attributes 12-15.
Edgecombe and Chatterton (1990) have discussed this character
and shown its plesiomorphic distribution; it is nonetheless not
constant within the taxa here considered.
2) Glabellar shape; 0 = longer than broad; 1 = subequal, within mea-
surement, preservation bias, or errors, as well as intrapopulation
variation; 2 = broader than long. This multistate character is con-
sidered unordered.
Edgecombe and Chatterton (1990) used this character, but did
not recognize state 1 above.
3) Maximum width of glabella; 0 = across the posterior part of gla-
bellar lobe L4; 1 = maximum width elsewhere.
If the maximum width is not across the posterior part of the
frontal lobe, it is anterior to that point.
4) Tubercles on anterior border of cephalon; 0 = absent; 1 = present.
This is essentially Temple and Tripp's (1979) attribute 28.
5) Number of axial rings in pygidium; 0 = 7-9; 1 = <7-9, > 7-9.
This continuous character is difficult to assess because of the
faintness of the posterior rings. Its importance, however, has been
discussed by Ramskold (1986, p. 529-531) who has suggested
alternate ways of coding the pygidial axis. Encrinurus hornei has
8 axial rings; in the natural sciences, values with an uncertainty
of ± 10% are the norm, which suggests that assigning state 0 to 7-
9 pygidial rings is reasonable, although it is uncertain if early
cybelines or encrinurines have less than 7 axial rings.
6) Number of congruent axial rings and pleural ribs; 0 = 4; 1 = <4,
>4.
This character has been defined by Temple and Tripp (1979) as
attribute 9 (see also Ramskold, 1986).
7) Number of pleural ribs; 0 = 5; 1 = ~6.
Although this character is generally coded as "pleural segments,"
"pleural ribs" are better terms, as ribs can be identified easily.
Whether these ribs correspond or not with pleurae is still open to
question (see Hessler, 1962, on this subject). In any event, axial
segmentation is poorly correlated with pleural architecture in en-
crinurines, which explains the usefulness ofcharacter 6. This char-
acter is the same as attribute 6 of Temple and Tripp (1979) and,
used somewhat differently, character 18 of Edgecombe and Chat-
terton (1990). These two authors have also shown the limited value
of this character within the encrinurines; as employed here, how-
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19
ever, this character is less informative than if it were treated as a
continuous variable.
8) Nature of genal spine; 0 = short (exsag. length shorter or approx-
'imately of the same (sag.) length as the occipital ring); 1 = longer
than state O.
This is Temple and Tripp's (1979) attribute 10, somewhat mod-
ified.
9) Lateral outline of glabella; 0 = straight; 1 = concave outwards; 2
= convex outwards. This multistate character is considered unor-
dered.
This is Temple and Tripp's (1979) attribute 32.
10) Ornamentation of occipital ring; 0 = smooth or granulose; 1 = with
one or more tubercles.
11) Adaxial tubercles on fixigenae; 0 = none; 1 = with one or more
tubercles.
This character is slightly modified from Temple and Tripp's
(1979) attribute 29.
12) Sagittal and nearly sagittal tubercles on axis of pygidium; 0 = with
at least one sagittal tubercle; 1 = with paired near sagittal tubercles;
2 = without tubercles. This multistate character is considered
unordered.
This is Edgecombe and Chatterton's (1990) character 4.
13) Axial thoracic spines; 0 = present on at least one segment;
without axial thoracic spines.
This is Edgecombe and Chatterton's (1990) character 11.
14) Postero-Iateral hypostomal spines in holaspis; 0 = absent; 1 = with
spines.
This is Edgecombe and Chatterton's (1990) character 6.
15) Glabellar ornamentation; 0 = uniform; 1 = not uniform.
16) Middle body of hypostoma; 0 = elongated; 1 = round (equally wide
as long).
The hypostomata of early encrinurines are inflated; this char-
acter is inspired by Edgecombe and Chatterton's comments (1990,
p. 823) on Physemataspis.
17) Pygidial shape; 0 = broader than long; 1 = subequal, within mea-
surement, preservation bias or errors, as well as intrapopulation
variation; 2 = longer than broad. This multistate character is con-
sidered unordered.
Temple and Tripp (1979) did not recognize posterior pygidial
spines (mucros) in the taxa treated here. State 2 of this character
does not occur in the early encrinurines considered here, but does
occur in the early cybelines of Table 1. The state of this character
for Frencrinuroides s.l. tholus n. gen. is discussed in the Systematic
Section.