Journal of Paleontology: Middle Cambrian (Acadian series) conoc...

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Middle Cambrian (Acadian series) conocoryphid and paradoxidid trilobites from the upper Chamberlain's Brook formation, Newfoundland and New Brunswick Kim, Dong Hee ABSTRACT-The Fossil Brook Member of the upper Chamberlain's Brook Formation is a thin (up to 14 m) but distinctive, unconformity-- bound depositional sequence recognizable from Rhode Island to eastern Newfoundland in Avalonian North America. Its diverse trilobite fauna was first described more than century ago from the limestone-rich facies of the member in southern New Brunswick. However, the systematics, stratigraphic context, and biostratigraphic significance of these trilobites have remained poorly known. A revision of the conocoryphid and paradoxidid trilobites has been completed, and the taxa set into their stratigraphic context within the middle Middle Cambrian. The faunas of the Fossil Brook are assigned to the Eccaparadoxides eteminicus Zone of Avalon. Although biogeographic barriers between Avalon and Gondwana remained strong in the Middle Cambrian and few shared trilobite species are present, a generalized correlation of the E. eteminicus Zone into Gondwana is with the Badulesia tenera Zone of the Toushamian Stage in Morocco and the Badulesia Zone of the Caesaraugustian Stage in Spain. INTRODUCTION CAMBRIAN SUCCESSIONS of the Avalon zone in the northeastern Appalachians and southern Britain are highly fossiliferous, well understood stratigraphically, and contain numerous volcanic ashes that can be precisely dated (see Landing, 1996). Consequently, they were not only appropriate for definition of the Precambrian-Cambrian boundary (Landing, 1994), but also offer the potential as a standard for global correlation and subdivision of the Cambrian. Unfortunately, limited information exists in earlier reports on the detailed biostratigraphy of the Avalonian trilobite-- bearing Cambrian and no data are generally available on the lithostratigraphic context and paleoenvironmental setting of the trilobites. Many species have never been illustrated photographically. This paper continues a revision of the Lower and Middle Cambrian trilobites of the Avalon zone in Appalachian North America (see Westrop and Landing, 2000; Geyer and Landing, 2001). Here we document the paradoxidids and conocoryphids of the Fossil Brook Member of the Chamberlain's Brook Formation, a thin but distinctive, unconformity-bound sequence that can be traced throughout Avalonian North America (Landing, 1996); agnostoid and solenopleuroid trilobites will be treated elsewhere. FOSSIL BROOK MEMBER Geologic setting.-The Fossil Brook Member is a lithologically distinctive, thin (up to 14 m) unit that extends from Rhode Island to east Newfoundland in the Avalon terrane of the northern Appalachians. Its stratigraphic and tectonic context has been clarified only recently (Landing, 1996; Landing and Westrop, 1998a, 1998b). The Fossil Brook is referred to depositional sequence 7 of the latest Precambrian-Lower Ordovician cool water succession of the Avalon continent (Landing, 1996). It forms the top of the green, purple, and red siliciclastic mudstone-dominated Chamberlain's Brook Formation (lower Middle Cambrian, up to 160 m thick) and underlies the dark gray and black mudstone-dominated Manuels River Formation (middle Middle Cambrian, up to 60 m thick). Significant basin reorganization in the transtensional tectonic regime of the Avalon continent controlled regional onlap and deposition, lateral facies changes, the regular appearance of trilobite-rich limestones (otherwise rare lower in the Chamberlain's Brook Formation), and development of the erosive lower and upper contacts of the Fossil Brook Member. East Newfoundland.-Episodes of volcanism and movements on growth faults bracketed Fossil Brook deposition in east Newfoundland. The greatest thicknesses of the Chamberlain's Brook Formation in this area lie along the elongate, fault-defined St. Mary's-east Trinity axis (Fig. 1), where volcanism took place just prior to Fossil Brook deposition. A 1.0-4.5 rn lens of calcitized volcaniclastic debris flows 4.5 rn below the Fossil Brook Member at Hopeall Head, east Trinity Bay (Hutchinson, 1962, p. 132, bed 28), and 60+ rn of pillow basalt in the underlying upper Braintree Member at Cape Dog, west St. Mary's Bay (Hutchinson, 1962; McCartney, 1967; Fig. 1) record volcanism shortly before deposition of the Fossil Brook. The top of the member across east Newfoundland is defined by a thin basaltic ash which separates the dark green-gray mudstones of the Fossil Brook from the slightly darker mudstone of the lower Manuels River Formation (Landing and Westrop, 1998b). The base of the Fossil Brook Member in east Newfoundland is a limestone with reworked shale and phosphatic granules that fill wide (up to 8 cm wide) Cruziana-like burrows deeply incised into overcompacted mud exposed by marine erosion (Landing, 1996, fig. 3, compare Landing and Brett, 1987). The thickness of the Fossil Brook ranges from 14 m at Deep Cove near the south end of the St. Mary's-east Trinity axis to 2.2-3.5 rn further north in Conception and Trinity bays and to less than a meter east in the Burin Peninsula. This reduction in thickness is attributed both to post-Fossil Brook erosion and stratigraphic condensation at northerly and easterly localities. Stratigraphic non-continuity and an upper sequence boundary are shown by a decrease in thickness from 3.4 m at Manuels River to 2.2 m at Red Bridge Road 8.5 km to the southwest and by erosional cut-out of several nodular limestone beds at the member's top (Landing and Westrop, 1998a, p. 27; Figs. 1, 2). Stratigraphic condensation is also suggested by the replacement of the siliciclastic mudstone-dominated Fossil Brook facies at Deep Cove (Fig. 1) and other localities in southern St. Mary's Bay by alternations of mudstone with wave-winnowed and sorted trilobite pack- and grainstones along Trinity and Conception bays. These fossil hash-rich, wave-influenced limestones are characteristic of higher-energy, near shore environments in Avalonian shale basins (e.g., Myrow and Landing, 1993). Maximum sedimentary condensation of the Fossil Brook member is seen at Little Dantzig Cove in the southwestern Burin Peninsula (Fig. 1), where the member comprises a 1.0 m bed of phosphatic limestone with numerous

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hardgrounds (Hutchinson, 1962, p. 146, bed 19). Nova Scotia.-Limited outcrop of the Chamberlain's Brook Formation precludes recognition of the Fossil Brook Member in southern Cape Breton Island and Antigonish area of mainland Nova Scotia (e.g., Landing, 1991, 1995; Fig. 1). Hutchinson (1952, p. 73, 74, pl. I, fig. 17) reported a cranidium of Paradoxides eteminicus, a characteristic form in the Fossil Brook in New Brunswick, from the east side of Mira Lake in Cape Breton Island. However, the specimen is tectonically distorted and cannot be identified to the species level. Examination of the dark shales in the vicinity of Hutchinson's locality suggests they are best referred to the younger Manuels River Formation (EL, unpub. data). New Brunswick.-The Fossil Brook reappears in its type area in the region of Saint John, New Brunswick, where a major unconformity separates it from the underlying upper Lower Cambrian Hanford Brook Formation and an upper unconformity with the Manuels River Formation can be demonstrated locally. Absence of the lower Chamberlain's Brook Formation (Braintree Member) and its lowest Middle Cambrian faunas (see Geyer and Landing, 2001) means that the Fossil Brook Member is the only remaining part of the formation in New Brunswick. The Fossil Brook overlies the Long Island Member (Kingaspidoides cf. obliquoculatus Zone) of the Hanford Brook Formation north and northeast of Saint John but rests on the older Somerset Street Member (Protolenus elegans Zone) in Saint John (Landing and Westrop, 1998a; Westrop and Landing, 2000). These data indicate as much as 45 m of differential erosion of the Hanford Brook Formation and suggest vertical movements on growth faults prior to marine onlap and Fossil Brook deposition. The unconformity at the base of the Fossil Brook Member is emphasized by a basal 30 cm black phosphatic quartz arenite with phosphatic pebbles that is overlain by 80 cm of nodular limestones with wave-comminuted trilobite hash and phosphate granules in Saint John (Figs. 1, 2, localities 113 and SoS). At other localities (GoS in Saint John, Caton's Island, and Hanford Brook West sections; Figs. 1, 2), the lenticular limestones in green-gray siltstone are the first Fossil Brook deposits above the non-calcareous, fine-grained sandstones of the Hanford Brook. Limestone lenses disappear above the lower several meters of the Fossil Brook, and the upper part of this thin unit (up to 13.3 m at Hanford Brook West) is dominated by laminated (non-burrowed) green-gray siliciclastic mudstones. As in east Newfoundland, the contact with the overlying Manuels River Formation is marked by an abrupt change to dark gray to black mudstones (Hayes and Howell, 1937; Landing, 1996), but a volcanic ash is not present. Evidence for a sequence boundary on the Fossil Brook Member comparable to that in east Newfoundland is now recognized at an outcrop at Caton's Island, north of Saint John (Fig. 1). There, a very thin Fossil Brook Member that consists of 1.3 m of limestone is directly overlain by black Manuels River Formation shales. Absence of the siltstones that comprise the upper Fossil Brook and presence of 20 cm of black phosphatic shale with reworked phosphatic granules at the base of the Manuels River are consistent with erosional beveling of the Fossil Brook. Rhode Island.-The Fossil Brook Member apparently exists in the structurally complex sections in southern Narragansett Bay, Rhode Island, where Skehan et al. (1978) recovered Badulesia tenera (Hartt in Dawson, 1868), a form known elsewhere in Avalon only in the Fossil Brook Member, from light gray phyllite. The upward succession from this phyllite into alternating light and dark gray phyllite, thin sandstones and shales, and finally dark gray phyllites with carbonate nodules suggests the Middle-Upper Cambrian succession (Fossil Brook-Manuels River Formation-MacLean Brook Group-Chesley Drive) elsewhere in Avalon (Landing, 1996). Earlier trilobite studies.-Most of the trilobites from the Fossil Brook Member were first described more than a century ago by C. E Hartt (in Dawson, 1868) and Matthew (1882-1885, 1890) from southern New Brunswick. Matthew (e.g., 1889) distinguished the limestones and gray mudstones of the member as a separate stratigraphic interval. He called it "division Ic" and recognized two successive "bands" or subzones, the lower with a Paradoxides lamellatus fauna (band lc') and an upper with a P. eteminicus fauna (Ic2). The composition of and differences between these faunas never seems to have been recorded, and the subzones seem to have been defined on order of appearance of the eponymous taxa. Hayes and Howell (1937, p. 69-72) qualified Matthew's faunal subdivision of the Fossil Brook by reporting that P. eteminicus ranged through the unit and preferred "P. lamellatus Zone" for the lower limestone and siltstones and "Hartella matthewi Zone" for the upper siltstones. Trilobite studies on the Middle Cambrian in eastern Newfoundland repeatedly focused on the Manuels River section (Fig. 1) and included recovery of trilobites through the Chamberlain's Brook Formation (e.g., Whiteaves, 1878; Howell, 1925). The latter author proposed generalized correlations of the entire Chamberlain's Brook in Newfoundland with "division Ic." However, Howell's (1925, p. 50-54) detailed collecting at Manuels showed that the Hartella, Bailiaspis, and Eccaparadoxides species (described below) characteristic of "division Ic" and the Fossil Brook member in New Brunswick abruptly appear in bedded limestones at the top of the Chamberlain's Brook beds 19-35 and do not persist into the Manuels River Formation. Howell's (1925) beds 19-35 at Manuels River are now referred to the Fossil Brook Member (Landing, 1996). Other than Paradoxides parvoculus Howell, 1925, which has yet to be encountered outside of the type locality (Hutchison, 1962), Howell did not illustrate any trilobites from the Fossil Brook Member in Newfoundland. Subsequent workers (e.g., Hutchinson, 1962; Martin and Dean, 1988) neither recognized the biostratigraphic importance of trilobites at the top of the Chamberlain's Brook Formation for precise correlation into New Brunswick nor undertook a comprehensive revision of the fauna. Newfoundland localities.-All localities of the Fossil Brook Member in eastern Newfoundland were examined. The top of the Chamberlain's Brook is most accessible in a road metal quarry located south of Killigrews village and 7.0 km east of the intersection of Red Bridge Road with coastal Rte. 60 (Landing and Westrop, 1998a, p. 18-22, figs. 4, 5). Exposures of gently west-- dipping uppermost Chamberlain's Brook form a bench below a highwall of lower Manuels River Formation immediately north of Red Bridge Road (Fig. 1). The Fossil Brook Member at Red Bridge Road (Fig. 2) is lithologically comparable to correlative strata 8.5 km to the northeast at Howell's (1925, beds 19-35) section at the Manuels River nature

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preserve. We found that Fossil Brook limestones on the east and west sides of Trinity Bay at Hopeall Head (Hutchinson, 1962, p. 132, beds 30-32) and Fosters Point on Trinity Island (Martin and Dean, 1988) were too indurated and tectonized, respectively, to yield material for taxonomic study. As noted above, the Fossil Brook is reduced to a single highly phosphatic limestone bed at Little Dantzig Cove and Snooks Brook in the Burin Peninsula (Fig. 1), and induration precluded recovery of specimens suitable for taxonomic study. New Brunswick localities.-The Fossil Brook Member overlies the Hanford Brook Formation at a number of localities described earlier by Landing and Westrop (1996, 1998a; Westrop and Landing, 2000). These include south-dipping exposures of the basal beds at three adjacent localities in Saint John: 113, a road ditch exposure on the north side of Exit 113 from Rte. 100; GoS, a low outcrop immediately west of the T-intersection of Seeley Street with Gooderich Street, and SoS, the basal conglomeratic bed of the Fossil Brook at the top of the road cut on the east side of Somerset Street (Fig. 1). All of the Fossil Brook is exposed in Glen Falls village just east of Saint John at locality GFW and on Hanford Brook at locality HBW (Fig. 1). GFW is on the south bank of Cold Brook downstream of the Glenview Avenue bridge. HBW (Hanford Brook West), on Hanford Brook about 14 km north of St. Martins village and just north of Rte. Ill (Landing and Westrop, 1998a, p. 61-65), has the lower 1.25 m of the Fossil Brook in the south cut-bank, with higher strata only visible at low water. Other localities northeast of Saint John include lower Fossil Brook Member sections on Ratcliffe Brook (locality RBr) just north of Quoddy Road and immediately downstream of the falls and Hayes and Howell's (1937) type locality on Fossil Brook (FoB-FB). FoB-FB is an exposure in the bed of Porter Road and immediately north in the Fossil Brook stream-cut (Landing and Westrop, 1998a, p. 65, 66). A thin, erosionally truncated section in the Fossil Brook is located north of Saint John on Caton's Island (locality CIE) in Saint John River (Fig. 1). CIE is exposed just on the middle part of the east shore of Caton's Island and immediately north of a low cliff of Hanford Brook Formation sandstone. Collateral collections.-Attempts were made to supplement our field work with museum collections. C. E Hartt's types were reposited at Cornell University in Ithaca, New York. They were later transferred to the Paleontological Research Institute in nearby Trumansburg, but cannot be located at the present time (P. Krohn, personal commun., September, 2000). The G. F Matthew Collection at the Royal Ontario Museum in Toronto includes topotype material of all of the species described by Hartt, and we have used them to supplement our field collections. An additional collection examined in this study was made at Seely Street, immediately east of our Gooderich Street locality, in Saint John, New Brunswick, by C. N. Hartnagel during the 1912 International Geological Congress and was found in the New York State Museum. AGE AND CORRELATION In New Brunswick, Matthew (1889) and Hayes and Howell (1937) noted that Eccaparadoxides lamellatus was restricted to the lower part of the Fossil Brook Member. Our collections do not contradict this observation, although we found E. lamellatus to be rare and recovered only a few cranidia in the basal limestone at Fossil Brook itself (FoB-FB 2.5; Fig. 2). Because of the paucity of data, we choose not to divide the Fossil Brook into two biostratigraphic units; rather, we assign the entire unit to the Eccaparadoxides eteminicus Zone. The eponymous species occurs throughout the member, as noted by Hayes and Howell (1937), including the basal bed at section 113 (113-14.85; Fig. 2). With the exception of an undescribed species of Bailiella and Eccaparadoxides cf. E. eteminicus, all of the other species treated here appear above the basal limestone of the Fossil Brook. Among the conocoryphids and paradoxidids, only the species of Eccaparadoxides (i.e., E. eteminicus, E. acadicus, and E. lamellatus) are shared between Newfoundland and New Brunswick. None of these species occur outside of Avalonian North America. However, the solenopleurid Badulesia tenera (C. E Hartt in Dawson, 1868) has been recorded from New Brunswick (Walcott, 1884; Kim, Westrop, and Landing, unpublished data), Newfoundland (Martin and Dean, 1988; Kim, Westrop, and Landing, unpublished data), Rhode Island (Skehan et al., 1978), Spain (Sdzuy, 1967), and Morocco (Sdzuy et al., 1999). These occurrences of B. tenera allow correlation of the Eccaparadoxides eteminicus Zone with the base of the Caesaraugustian Stage in Spain and middle of the Toushamian Stage in Morocco, and can be informally regarded as indicating an interval in the transition from the lower Middle to middle Middle Cambrian (see also Geyer, 1998, fig. 2; Sdzuy et al., 1999, fig. 5). SYSTEMATIC PALEONTOLOGY Trilobite systematics should be attributed to Kim and Westrop in Kim et al.; the order of these names is alphabetical and does not indicate seniority. Illustrated material is housed in the Royal Ontario Museum (ROM), the New Brunswick Museum (NBMG), and the New York State Museum (NYSM). ROM material designated as "referred specimens" in the text are from the G. E Matthew collection and were identified, but not figured, by Matthew. Superfamily SOLENOPLEUROIDEA Angelin, 1854 Family CONOCORYPHIDAE Angelin, 1854, emend. Cotton, 2001 Discussion.-Several workers (e.g., Fortey, 1990; Geyer, 1998) have suggested that the traditional concept (e.g., Poulsen, 1959) of the Conocoryphidae may be an artificial grouping of taxa that lack eyes and have marginal or submarginal sutures. In a recent phylogenetic analysis, Cotton (2001) confirmed this suspicion by demonstrating that the family is polyphyletic and comprises four distantly related Glades. He also showed that a monophyletic Conocoryphidae should be restricted to Conocoryphe Hawle and Corda, 1847, Ctenocephalus Hawle and

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Corda, 1847, Elyx Angelin, 1854, Holocephalina Salter, 1864, Bailiella Matthew, 1885, Hartella Matthew, 1885, Cainatops Matthew, 1889, Bailiaspis Resser, 1936, Parabailiella Thoral, 1946, and Tchaiaspis Korobov, 1966. This emended concept of the family will be followed herein, along with Cotton's (2001, p. 192) revised diagnosis. Ingroup relationships of the conocoryphid genera were poorly resolved in Cotton's analysis and several species were unstable if different outgroups were used to root the cladogram (Cotton, 2001, fig. 4). As recognized by Cotton, resolution of these problems will require a better understanding of the sister groups of the Conocoryphidae among the "ptychoparioids." It will also require a more comprehensive analysis of the full compliment of valid conocoryphid species instead of the subset used by Cotton. Such an analysis is beyond the scope of this paper, but preliminary comments on some conocoryphid genera will be made below. Genus BAILIELLA Matthew, 1885 Type species.Conocephalites baileyi C. F Hartt in Dawson, 1868 from the Chamberlain's Brook Formation of New Brunswick, Canada (subsequent designation by Miller, 1889). Discussion.-Bailiaspis Resser, 1936, was established for species that differed from Bailiella baileyi by possessing a plectrum (e.g., Fig. 4.1-4.4; Walcott, 1884, pl. 4, fig. 2, 2b; Westergard, 1950, pl. 6, figs. 4, 5; Egorova et al., 1982, pl. 1, fig. 8, pl. 11, fig. 11). However, in lacking a plectrum, the border of Bailiella (Figs. 3.1, 3.4, 4.7) is similar to those of other conocoryphid genera, including Conocoryphe itself (e.g., Snajdr, 1958, pl. 31, figs. 8-13; Hutchinson, 1962, pl. 13, fig. 8a), and is most likely plesiomorphic. Where known from uncompacted material, Bailiella possesses weakly inflated fixigenae (e.g., Westergard, 1950, pl. 5, figs. 5b, 6b, 6c, 10c), but the same character state is also present in Conocoryphe (e.g., Hutchinson, 1962, pl. 13, fig. 7b) and Elyx (e.g., Hutchinson, 1962, p1. 12, fig. 5b; Westergard, 1950, pl. 7, fig. 3c), and may be plesiomorphic. Many conocoryphids possess sutures that are marginal, or very nearly so, so that at least part of the border and border furrow can be traced along the lateral margin to the level of the Os furrow (e.g., Westergard, 1950, pi. 6, fig. 9a, p1. 7, figs. 1-4; Hutchinson, 1962, p1. 10, fig. 18a; Dean, 1982, fig. 11). In contrast, the sutures of Bailiella swing inward at, or in front of, glabella mid-length to truncate the lateral border and border furrow (e.g., Fig. 3.1; West(egard, 1950, p. 24, pl. 5, figs. 1-3, 6-8, 10-12, pl. 6, figs. 1-3; Snajdr, 1958, pl. 35, fig. 1; Geyer, 1998, pl. 4, fig. 4). However, this state also occurs in Bailiaspis (e.g., Westergard, 1950, p1. 6, figs. 4, 5) and in some species of Conocoryphe (e.g., Sdzuy, 1967, p1. 10, figs. 7-15, 18-21, 23) and does not provide unequivocal support for monophyly of Bailiella. Moreover, some species currently assigned to Bailiella (e.g., Zhang and Jell, 1987, p1. 35, figs. 1-7) possess lateral borders that extend back to join the posterior border. In the absence of unambiguous apomorphies, Bailiella is probably paraphyletic and no attempt will be made to frame a diagnosis. Cotton's (2001) parsimony analysis of the Conocoryphidae also failed to find compelling support for monophyly of Bailiella, and the genus is used herein with reservation. BAILELLA BAILEYI (Hartt in Dawson, 1868) Figure 4.6-4.8 Conocephalites baileyi HAR-r in DAWSON, 1868, p. 645 Conocorype (Bailiella) baileyi (Hartt in Dawson). MATTHEW, 1885, p. 111, figs. 22-29. Conocoryphe (Salteria) baileyi (Hartt in Dawson). WALCOT, 1884, p. 312, pl. 4, fig. 3, 3a, pl. 5, fig. 7, 7a. Conocoryphe baileyi (Hartt in Dawson). MATTHEw, 1890, p. 135, p1. 11, fig. 10. Bailiella baileyi (Hartt in Dawson). RESSER, 1936, p. 16; SHIMER AND SHROCK, 1944, p. 607, pl. 253, figs. 10-12; COTTON, 2001, text-fig. lB, pi. 4, figs. 2-3. Diagnosis.-A species of Bailiella with cranidial internal molds smooth except for caecal network associated with prominent palpebral ridge. Long preglabellar field equal to about twice anterior border length. Transversely subelliptical pygidium with axis of two well-defined rings plus terminal piece. Description.--Cranidium semielliptical in outline, with width greater than length. Anteriorly rounded glabella tapers forward and occupies about 70 percent of cranidial length; axial and preglabellar furrows firmly impressed (possibly exaggerated by compaction). Barely perceptible, oblique SI and S2 lateral furrows present on some specimens; occipital furrow shallow and subtransverse medially, but curves forward distally. Long preglabellar field equal to about twice anterior border length; anterior border furrow shallow and curved forward. Fixigenae broad, wider than glabella at posterior. Palpebral ridges originate at subcircular caecal masses near anterior end of glabella, curving backward across cheek to terminate opposite glabellar mid-length; caecal networks originate from ridges and are best developed anteriorly. Broad, shallow, posterior border furrow defines narrow posterior border. Facial suture truncates border and border furrow opposite S2 glabellar furrow. Apart from caecal networks, internal molds are smooth. Pygidium is transversely subelliptical in outline, with convex axis that occupies about 80 percent of pygidial length. Two axial rings and terminal piece present; deep articulating furrow defines short articulating half-ring, but axial and axial ring furrows shallow. Three pleural

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furrows cross broad pleural field, with anteriormost much deeper than others; interpleural and border furrows obsolete. Surface of internal mold is smooth. Material examined.--Six cranidia and two pygidia from the Matthew collection, ROM, collected at Seely Street, Saint John; five cranidia from NYSM, also from Seely Street. Discussion.-Bailiella baileyi (Hartt, in Dawson, 1868), although originally described from New Brunswick, was not encountered in any of our collections and is illustrated with topotype material from the Matthew collection at the Royal Ontario Museum. All specimens have been deformed to varying extents by compaction, so that such features as convexity of various elements of the exoskeleton have not been included in the description. Compaction also hinders comparisons with other species. Some, including Bailiella tenuicincta (Linnarsson; Westergard, 1950, p1. 5, figs. 6-8) and B. aequalis (Linnarsson; Westergard, 1950, p1. 5, fig. 13) differ from B. baileyi by possession of granulose or tuberculate sculpture. Bailiella manuelensis (Fig. 3.1, 3.2), from Newfoundland, and B. meridiana (Sdzuy; Sdzuy, 1961, pi. 34, fig. 5), from Spain, differ from B. baileyi by possession of a very short preglabellar field. Bailiella pokrovskavae Koborov (1973, Egorova et al., 1976, pi. 35, figs. 11-13), from Siberia, and B. lantenoisi Mansuy (Zhang and Jell, 1987, pl. 35, figs. 1-7) both resemble B. baileyi in the length of the preglabellar field, but are differentiated easily by the length of the border. In B. pokrovskayae, the short anterior border is truncated by the facial sutures in front of the anterior end of the glabella, whereas the border extends back to join the posterior border in B. lantenoisi. Among species from Avalonian Britain, B. comleyensis Resser (Lake, 1940, pl. 60, figs. 7, 8), B. longifrons (Cobbold; Lake, 1940, pi. 60, figs. 9-11) and B. cobboldi Resser (Lake, 1940, p1. 60, figs. 12-14) all possess narrower cranidia than B. baileyi, in which the posterior widths of fixigena are roughly equal to the widths of the glabellae. Bailiella lyelli (Hicks; Lake, 1940, p1. 60, figs. 1-6) is based on deformed material, so that comparisons are difficult. Cranidia of B. lvelli appear to possess deeper lateral glabellar furrows and conspicuous, convex palpebral ridges, whereas the pygidium appears to be relatively shorter and narrower. Bailiella barriensis Sdzuy (1958; 1961, pi. 33, figs. 1-5), from the Middle Cambrian of Spain, and B. howelli (Hutchinson, 1962, p1. 14, figs. 1-4), from Newfoundland possess anterior borders that are longer (sag) than that of B. baileyi, and both lack palpebral ridges. The Moroccan species, B. inconspicua Geyer (1998, pl. 4, figs. 1-3) and B. dilatata Geyer (1998, pl. 4, figs. 4, 7) are also characterized by much longer anterior borders which, in the case of the former, is upturned. BAILIELLA WALCOTTI (Matthew, 1885) Figure 3.3-3.8 Conocoryphe (Bailiella) walcotti MATTHEW, 1885, p. 119, fig. 36, 366. Conocoryphe walcotti (Matthew). WALCOTT, 1884, p. 310; MATTHEW, 1890, p. 134, pl. 11, fig. 7a-c. Bailiella walcotti (Matthew). RESSER, 1936, p. 16. Diagnosis.-A species of Bailiella with broad cranidium with sculpture of scattered fine granules on internal molds. Pygidial axis long, with three rings and terminal piece. Type.-Lectotype, cranidium (ROM 54058) from the Fossil Brook Member, Chamberlain's Brook Formation of New Brunswick (Fig. 3.4), selected herein. Material examined.-Forty-seven cranidia and seventeen pygidia from field collections plus eight cranidia from Matthew's type lot. Occurrence.-Fossil Brook Member, New Brunswick, collections FoB-FB 0.2, 2.5; GoS 2.3; 113-14.85; Matthew's types are from Saint John, most likely Seely Street. Discussion.-Bailiella walcotti Matthew is very similar to B. baileyi (Hartt), so that a comparison is presented instead of a full description. Cranidia of B. walcotti differ only by possession of sculpture of fine granules on internal molds (Fig. 3.4, 3.8) that becomes more subdued on larger holaspids (Fig. 3.3) The pygidium of B. walcotti was previously unknown. It differs from that of B. baile.vi (Fig. 4.6) in having a longer axis with three, rather than two, axial rings plus terminal piece; pleural furrows are more firmly impressed. The pygidium of B. baileyi is relatively shorter and wider, although the difference in outline may have been exaggerated by compaction. Most of the differences that separate B. baileyi from other species (discussed above) apply equally to B. walcotti. Like B. walcotti, B. tenuicincta (Linnarsson; Westergard, 1950, pl. 5, figs. 68) and B. aequalis (Linnarsson; Westergard, 1950, pl. 5, fig. 13; and its possible synonym, B. ornata Resser, 1937) possess granulose or tuberculate sculpture. In B. aequalis, the granules are larger and closely spaced but, in general cranidial proportions, this species is close to B. walcotti. Sculpture is comparable in B. tenuicincta, but the cranidium is narrower, and the pygidium consists of only two rings and a terminal piece. BAILIELLA MANUELENSIS Hutchinson, 1962 Figure 3.1, 3.2

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Bailiella manuelensis HUTCHINSON, 1962, P. 106-107, pi. 15, fig: 13; LwSs. 5-; D,in KAILinN Am DEAN, 1988, P. 19, pl. 2, fig. 13; Lim-Sum, 1993, p1. 71. Diagnosis.-A species of Bailiella with subrectangular cranidium with weakly curved anterior margin. Short preglabellar field equal to about 12 percent of glabellar length. Strongly tapered, conical glabella with barely perceptible lateral furrows. Surface of cranidum smooth except for faint palpebral ridge and associated caecal network. Material examined.-Four complete exoskeletons and five cranidia. Occurrence.-Fossil Brook Member, southeastern Newfoundland, collection RBCB 49.5-50.2. Discussion.-There is little to be added to Hutchinson's (1962, p. 106-107) original description of Bailiella manuelensis. A well preserved, nearly complete exoskeleton (Fig. 3.1) shows that the tips of thoracic pleurae are not evenly rounded, but terminate at very short, bluntly-pointed spines. The short preglabellar field of Bailiella manuelensis separates it from a variety of species including B. lyelli (Hicks; Lake, 1940, pl. 60, fig. 6) and B. longifrons (Cobbold; Lake, 1940, pi. 60, fig. 9) from Avalonian Britain, B. tenuicincta (Linnarsson; WestergArd, 1950, pl. 5, figs. 6, 8) and B. impressa (Linnarsson; Westergard, 1950, p1. 5, figs. 10-12) from Sweden, B. dilatata Geyer (1998, pl. 4, figs. 4, 7) from Morocco, B. pokrovskayae Koborov (Egorova et al., 1976, pl. 35, figs. 11-13), from Siberia, and B. barriensis Sdzuy (1958; 1961, pl. 33, figs. 1-5), from Spain. Bailiella aequalis (Linnarsson; Westergard, 1950, pl. 6, figs. 1-3) and B. emarginata (Linnarsson; Westergard, 1950, pl. 5, figs. 1-4), from the Middle Cambrian of Sweden, have short preglabellar fields that approach B. manuelensis. Both of the Swedish species have shallow but clearly defined lateral glabellar furrows, and granulose sculpture on the cranidium. In addition, B. emarginata has a strongly curved anterior cranidial margin and fifteen, rather than fourteen, thoracic segments. BAILIELLA sp. Figure 4.9. 4.10 Material examined.-our cranidia. Occurrence.-Fossil Brook Member, New Brunswick, collections FoB-FB 0.2; 113-14.85. Discussion.-Bailiella sp. is unusual in two respects: the lateral border extends back along the cranidial margin to join the posterior border, and the sculpture consists of low rounded tubercles among a background of finer, more closely spaced granules. The most similar species is B. lantenoisi (Mansuy; Zhang and Jell, 1987, pl. 35, figs. 1-7), which differs in sculpture and in having a less convex border that is separated from the fixigenae only by a break in slope. In contrast, Bailiella sp. is characterized by deep border furrows. Genus BAILIASPIS Resser, 1936 Type species.-Conocephalites elegans Hartt in Dawson, 1868, from the Middle Cambrian of New Brunswick (by original designation). Discussion.-Westergard (1950, p. 24) noted that the medial thickening of the anterior border used by Resser (1936, 1937) to diagnose Bailiaspis could be produced in two ways. In some species, including the type species, Bailiaspis elegans (Hartt; Fig. 6.9, 6.10; Walcott, 1884, p1. 4, fig. 2, 2b). the border "expands backwards" (i.e., has a true plectrum), and the anterior border furrow is bowed sharply posteriorly. In other species, such as Conocoo,phe emarginata Linnarsson (WestergArd, 1950, pi. 5, figs. 1-4; assigned to Bailiaspis by Resser, 1936, p. 19), medial thickening reflects the course of the submarginal suture, and the anterior border furrow is not deflected backwards (i.e., a plectrum is absent). Such thickening occurs in other conocoryphid genera, including Conocoryphe (e.g., Dean, 1982, fig. 11a). We agree with Westergard (1950) that Bailiaspis should be restricted to species that possess a plectrum and, in addition to the species that he excluded, we transfer Bailiaspis howelli Hutchinson (1962, pl. 14, figs. 1-4) B. cf. howelli Hutchinson (1962, pl. 14, figs. 5-6) and B. meridiana Sdzuy (1958; Sdzuy, 1961, pl. 34, fig. 5) to Bailiella. As used herein, Bailiaspis includes, at minimum, B. elegans (Hartt in Dawson, 1868), B. dalmani (Angelin, 1878), B. venusta Resser, 1937 (=B. prominens Resser, 1937), B. inflata Lake, 1940, B. tuberculata Lake, 1940, B. latigenae Hutchinson, 1962, B. menneri Koborov, 1973, B. bobrovi Koborov, 1973, B. jakutensis Koborov, 1973, B. botomensis Koborov, 1973, B. picta Koborov, 1973, B. curta, Koborov, 1973, B. senonta, Koborov, 1973, and, possibly, B. glabrata (Angelin, 1878). Bailiaspis nicholasi Resser, in Lake (1940, pl. 61, figs. 1-4) is correctly assigned to the genus but is based on very poorly preserved, deformed material, and is best restricted to the types. Lake (1940, p. 283) suggested that Bailiaspis was differentiated from Bailiella by "depression of the preglabellar area so that the fixed cheeks are separated from each other instead of continuous." The depression actually reflects the fact that the fixigenae are inflated in Bailiaspis (Fig. 4.2) and, as used here, the genus comprises species with moderately to strongly inflated fixigenae. It is unclear as to whether Bailiaspis is monophyletic because a plectrum is also present in the poorly known genus Tchaiaspis Koborov, 1966. The plectrum of the type species, T. sdzuyi Koborov (Cotton, 2001, pl. 4, fig. 9), is unusually long and narrow, but is shorter and similar to the condition in Bailiaspis in the only other known species, T. sp. nov. St John and Babcock (1997, fig. 6R-U). Tchaiaspis is distinctive in possessing a well-defined paradoublural band at the inner edge of the lateral border furrow. The relationship between Tchaiaspis and Bailiaspis is not resolved clearly in Cotton's (2001) phylogenetic analysis, but some of his cladograms (2001, fig. 4) indicate that recognition of the former may create paraphyly in the latter. Thus, it may prove preferable to treat Tchaiaspis as a junior synonym of Bailiaspis.

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BAILIASPIS VENus-rA Resser, 1937 Figure 4.1-4.5 Bailiaspis venusta RESSER, 1937, p. 40, pl. 7, figs. 24-25; SHIMER AND SHROCK, 1944, p. 607, pi. 253, fig. 19; HUTCHINSON, 1962, p. 100, pl. 13, figs. 11-12. Bailiaspis prominens RESSER, 1937, p. 41, pl. 7, figs. 5, 6; HUTCHINSON, 1962, p. 101, pl. 13, figs. 13, 14 (see for synonymy). non Bailiaspis venusta RESSER, 1937. COTTON, 2001, pl. 4, fig. 7 [=Parasolenopleura? applanata (Salter in SALTER AND HICKS, 1869)]. Diagnosis.-A species of Bailiaspis with preglabellar field very short or absent. Occipital ring lacks occipital spine. Cranidial sculpture includes numerous densely-packed tubercles. Pygidium with broad, parallel-sided axis. Description.Cranidium convex and semielliptical in outline, with length about half of width. Forwardly tapered, bluntly rounded glabella with width at anterior about 40 percent of width at occipital furrow; axial furrows well incised and preglabellar furrow somewhat shallower. Three pairs of shallow lateral glabellar furrows present: SI long, directed obliquely backwards and extending a third across glabella; S2 short and oblique; S3 very short and subtransverse. Occipital furrow shallow, subtransverse medially, but curved forward distally. Occipital ring occupies about 20 percent of glabellar length and has gently curved posterior margin. Median portion of ring incomplete on all available specimens but apparently lacks occipital spine (Fig. 4.1; see also Hutchison, 1962, p. 101). Frontal area occupies about 25 percent of cranidial length. Preglabellar field very short or absent, with anterior border furrow and preglabellar furrow confluent. Anterior border convex with short plectrum, and narrows distally; anterior border furrow shallow but clearly defined and curved gently forward on either side of plectrum. Fixigenae convex, with crests below level of glabella; barely perceptible palpebral ridges on some specimens. Posterior border furrow is broad, shallow groove; posterior border narrow near axial furrow but widens distally. Facial sutures intersect border and border furrow opposite anterior end of glabella and diverge gently backwards to posterior corners of cranidium. Sculpture on all surfaces except furrows comprises closely spaced granules with more widely spaced tubercles. Pygidium moderately convex, semielliptical in outline, with length about half of width. Axis parallel-sided and rounded posteriorly, divided into two axial rings and a terminal piece. Axial and axial ring furrows very shallow. Pleural field weakly convex and slopes downward distally; border absent. Two pairs of pleural furrows, with anteriormost deeply incised; one pair of faint interpleural furrows present. Pleural bands and axial rings carry row of granules; remainder of surface smooth. Material examined.-Nine cranidia and one pygidium. Occurrence.-Fossil Brook Member, southeastern Newfoundland, collections RBCB 49.5-50.2. Discussion.-Hutchinson (1962, p. 101) recognized that Bailiaspis venusta Resser and B. prominens (Resser) were very similar species, and separated them only because of apparent differences in stratigraphic occurrence. Our cranidia (Fig. 4.1-4.4), collected from a stratigraphic interval of less than a meter at Red Bridge Road, show variability in features used by Resser (1937, p. 41) to differentiate these species, including glabellar outline, expression of glabellar furrows, and convexity of the anterior border. Expression of tuberculate sculpture is related to preservation and varies even within a single specimen (Fig. 4.3). For these reasons, we consider B. prominens to be a junior synonym of B. venusta. Cotton (2001, pl. 4, fig. 7) assigned a cranidium from the Manuels River Formation to B. venusta, but this species has been recorded only from the Chamberlain's Brook Formation by previous workers (Fig. 2; Hutchison, 1962). Cotton's specimen appears to possess a palpebral lobe on the left side and is transferred to Parasolenopleura? applanata (Salter in Salter and Hicks, 1869), a species that does occur in the Manuels River Formation (Martin and Dean, 1988, pl. 3, figs. 3, 6, 8, 14, 15). Bailiaspis elegans (Hartt; Walcott, 1884, pl. 4, fig. 2, 2b), from New Brunswick possesses a short occipital spine that is not present in B. venusta (see following discussion of B. elegans). Bailiaspis dalmani (Angelin; Westergard, 1950, p1. 6, figs. 4, 5) from Sweden, and the Siberian species B. jakutensis Koborov (Egorova, 1982, pl. 1, fig. 8, pl. 58, fig. 6), B. bobrovi Koborov (Egorova, 1982, pl. 11, fig. 11, pl. 16, figs. 9, 10), B. botomensis Koborov (1973, pl. 11, fig. 1), B. picta Koborov (1973, pl. 11, figs. 2, 3), B. curta Koborov (1973, pl. 11, fig. 4) and B. senonta Koborov (1973, pl. 11, fig. 6) have longer preglabellar fields than B. venusta, and less inflated fixigenae; B. in,/tata Lake (1940, pl. 61, figs. 5-7), from the Middle Cambrian of England, also possesses a longer preglabellar field and has a glabella that is nearly parallel-sided rather than conspicuously tapered. Bailiaspis latagenae Hutchinson (1962, pl. 14, fig. 7a, b), from the Manuels River Formation of Newfoundland, is separated readily from B. venusta by its much wider and more strongly inflated fixigenae, whereas B. tuberculata Lake from the Middle Cambrian of England, is characterized by a cranidial sculpture of closely-spaced, coarse tubercles. Finally, Bailiaspis menneri Koborov (1973, pl. 9, figs. 1-8) lacks a preglabellar field, so that the plectrum and glabella are separated only by the preglabellar furrow. BAILIASPIS ELEGANS (Hartt in Dawson, 1868) Figure 6.8-6.10 Conocephalites elegans HART in DAWSON, 1868, p. 650. Conocoryphe elegans (Hartt). WALCOTT, 1884, p. 313, pl. 4, figs. 2, 2b [only; Fig. 2a = Conocoryphe granulatus (Walcott)].

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Bailiaspis elegans (Hartt). RESSER, 1936, p. 18. Diagnosis.-A species of Bailiaspis with short occipital spine. Cranidial sculpture of tubercles scattered among densely packed granules. Pygidium with narrow axis of two rings and terminal piece; pleural furrows firmly impressed. Material examined.-Six cranidia and one pygidium from the Matthew Collection, ROM. Occurrence.-Fossil Brook Member, New Brunswick, Seely Street, Saint John, and St. Martin. Discussion.-Bailiaspis elegans was not encountered in our collections and is illustrated using referred specimens from the Royal Ontario Museum (Fig. 6.8-6.10). The cranidium is very similar to Bailiaspis venusta Resser (Fig. 4.1-4.4), so that a detailed description is not necessary. The occipital spine, illustrated clearly by Walcott (1884, fig. 2, 2b), is not preserved completely in any of our specimens but remains one of the few features that separates B. elegans from B. venusta. Although the latter lacks a spine, it may possess a small node (Hutchinson, 1962, p. 101). The only other difference between cranidia lies in the sculpture. Bailiaspis elegans (Fig. 6.9, 6.10) consistently displays tubercles scattered among fine granules, whereas the tubercles are much more densely packed on the surface of B. venusta (Fig. 4.1-4.4). A pygidium among the referred specimens of B. elegans (Fig. 6.8) differs from B. venusta (Fig. 4.5) in having a narrower, gently tapered axis and three pairs of well-defined pleural furrows. Genus CTENOCEPHALUS Hawle and Corda, 1847 Type species.--Ctenocephalus barrandei Hawle and Corda, 1847 [ = Conocephalus coronatus Barrande, 1846] from the Skryje Formation, Czech Republic (by original designation). Diagnosis.-A genus of Conocoryphidae with subcircular preglabellar boss separated from anterior border by well-defined border furrow. Discussion.One of the most stable parts of the recent analysis by Cotton (2001, figs. 2-4) dealt with the relationships of Ctenocephalus Hawle and Corda, Hartella Matthew and Elyx Angelin, three genera that possess a subcircular preglabellar boss. Monophyly of both Ctenocephalus and Elyx was supported by the analysis, but Hartella emerged as a paraphyletic grouping of species related to Ctenocephalus. Ctenocephalus as conceived by most recent authors (e.g., Hutchinson, 1962; DEAN in Martin and Dean, 1988) may be diagnosed by possession of a well-defined ridge along the fixigenae (e.g., Hutchinson, 1962, pl. 12, figs. 18a, 19a; Snajdr, 1958, pl. 36, figs. 1, 3, 7). By comparison with Hartella (e.g., Figs. 5.15.2, 5.5-5.10, 6.1-6.7), Elyx (e.g., Westergard, 1950, pl. 7, figs. 1-4) and other conocoryphids (e.g., Figs. 3.1-3.4, 4.1-4.4, 4.74.10, 6.10), this state is apomorphic and supports monophyly of Ctenocephalus. Elyx may be diagnosed by such apomorphies as fusion of the preglabellar boss to the anterior border (e.g., Westergard, 1950, pl. 7, fig. 4b) and a subrectangular to suboval cranidial outline (see also Hutchinson, 1962, pl. 13, fig. 4a; Babcock, 1994, fig. 8.1, 8.3a; Egorova et al., 1982, pl. 22, fig. 2). Where recognized as a separate taxon, Hartella has been diagnosed by the absence of a fixigenal ridge (e.g., Hutchinson, 1962; Dean in Martin and Dean, 1988), but this state occurs in all other conocoryphids, and is therefore plesiomorphic. The majority of sclerites of Hartella from Newfoundland (e.g., Fig. 6.4) and New Brunswick (e.g., Fig. 5.8) have undergone compaction, but specimens that retain most of their original convexity (e.g., Figs. 5.6, 5.7, 6.5, 6.6) demonstrate that the fixigenae were strongly inflated. Although this differs from the less convex condition in Ctenocephalus (Hutchinson, 1962, pl. 12, figs. 18b, 19b; pl. 13, fig. lb) Elyx (Westergard, 1950, pl. 7, fig. 3a, 3c) and most other conocoryphids (e.g., Westergard, 1950, pl. 5, figs. 5c, 8b, 10c, pl. 6, figs. 5c, 7c), fixigenal inflation does not appear to be an unambiguous synapomorphy for Hartella. Some species of Bailiaspis (e.g., Hutchinson, 1962, p. 103, pl. 14, fig. 7a, b) have strongly convex genae that stand above the level of the glabella, whereas some species of Hartella (e.g., Westergard, 1950, pl. 6, fig. 9c) have genae that are less convex than the glabella. Thus, there does not appear to be any character support for monophyly of Hartella and, from the structure of Cotton's (2001, figs. 2-4) cladograms, we conclude that the genus is best treated as a junior synonym of Ctenocephalus. This constitutes a return to the classification of Westergard (1950). As used herein, Ctenocephalus is a genus of Conocoryphidae in which a subcircular boss is separated from the anterior border by a well-defined border furrow. CTENOCEPHALUS MAT HEm (Hartt in Dawson, 1868) Figure 5.1-5.10 Conocephalites matthewi HART-r in DAWSON, 1868, p. 646, fig. 224. Conocephalites gemini-spinosus HARTT in DAWSON, 1868, p. 653. Conocoryphe matthewi (Hartt). WALCOTT.carr, 1884, p. 308, pl. 4, fig. 1, la, b. Ctenocephalus (Hartella) matthewi (Hartt). MATTHEW, 1885, p. 103, figs. 6-21; COTTON, 2001, p1. 4, figs. 10, 11. Ctenocephalus (Hartella) matthewi (Hartt). var. geminispinosus MATTHEW, 1885, p. 106. Ctenocephalus (Hartella) matthewi (Hartt). var. hispidus MATTHEW, 1885, p. 106.

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Ctenocephalus (Hartella) matthewi (Hartt). var. perhispidus MATTHEW, 1885, p. 107. Ctenocephalus matthewi (Hartt). RESSER, 1936, p. 20. Ctenocephalus hispidus (Matthew). RESSER, 1936, p. 20. Ctenocephalus perhispidus (Matthew). RESSER, 1936, p. 20. Ctenocephalus hispidus (Matthew). RESSER, 1936, p. 20. Ctenocephalus hartti RESSER, 1936, p. 20. Hartella matthewi (Hartt). DEAN in MARTIN AND DEAN, 1988, pi. 2, fig. 3. Diagnosis.-A species of Ctenocephalus that lacks fixigenal ridges and occipital spine. Glabella subtrapezoidal in outline. Lateral border of even width; lateral border furrow does not join posterior border furrow. Internal molds of cranidia have sculpture of fine granules with scattered, large tubercles, and palpebral ridges in form of caecal network. Description.-Cranidium strongly convex and senuelliptical in outline. Convex glabella subtrapezoidal in outline and occupies about 60 percent of cranidial length; axial furrows deeply incised and preglabellar furrow somewhat shallower but distinct. Three pairs of firmly impressed lateral glabellar furrows: SI oblique; S2 oblique and shorter than SI ; S3 very short and transverse. Occipital furrow wellincised, subtransverse medially but curves forward distally. Occipital ring occupies about 20 percent of glabellar length and carries median tubercle. Frontal area with strongly convex, subcircular preglabellar boss separated from short, convex anterior border by shallow, forwardly convex border furrow. Border furrow deepens away from boss; rim-like lateral border maintains even width back to border furrow. Broad fixigenae strongly convex, standing above level of glabella in least-compacted cranidia. On internal molds, weak palpebral ridges curve backward from anterior ends of glabella, terminating opposite L2 glabellar lobe; caecal network extends forward from palpebral ridge but is absent posteriorly. Posterior border narrow, convex rim near glabella, but widens somewhat distally; posterior border furrow well-defined and wide. Sculpture of scattered tubercles on glabella, boss, genae and borders; well-preserved specimens (Fig. 5.8) show that tubercles are interspersed among more densely-packed, granules. A poorly preserved specimen (Fig. 5.3) demonstrates that narrow librigena carries long genal spine. Pygidium sub-elliptical in outline, with strongly-curved anterior margin and gently curved posterior margin. Convex axis occupies about 33 percent of pygidial width and consists of single axial ring and terminal piece; long articulating half-ring at anterior end of axis. Axis occupies about 75 percent of pygidial length, and post-axial ridge extends to posterior margin. Pleurae crossed by single, shallow pleural furrow. Internal mold smooth. Material examined.-Thirty-three cranidia from field collections; five cranidia and one librigena from the Matthew Collection, ROM; ten cranidia from NYSM. Occurrence.-Fossil Brook Member, New Brunswick, collection FoB-FB 2.5; GoS 2.3.; Matthew's material at the ROM was collected from Seely Street, Saint John; NYSM material is from Seely Street and from Ratcliffe Brook (RBr). Discussion.-Resser (1936) elevated three varieties of C. matthewi recognized by Matthew (1885), C. hispida, C. perhispida and C. geminispinosa, to distinct species, and named a fourth species, C. hartti, for material illustrated by Walcott (1884, pl. 4, fig. 1). These differ minimally from C. matthewi and are regarded herein as synonyms. Resser (1936) also recognized Ctenocephalus matthewi var. granulatus Walcott (1884) as a distinct species. It is know from only two specimens from the Hartt collection (Walcott, 1884, pi. 4, figs. lb, 2a), both of which appear to have undergone considerable compaction. According to Walcott (1884, p. 310), the cranidium possesses closely spaced tubercles and "the occular-like ridges are lost in the crowding together of the tubercles." In these respects, it resembles C. terranovicus (Resser) from Newfoundland (see below). No cranidia with sculpture of the type described by Walcott are present in our field collections and, until Hartt's specimens can be found and examined, the status of C. granulatus is uncertain. Hutchinson (1962, p. 96) noted that there are few differences between Ctenocephalus matthewi and C. terranovicus (Fig. 6.16.7) that cannot be interpreted as a consequence of the degree of compaction. As he recognized, these very similar species are separable on the basis of sculpture and the presence of palpebral ridges on internal molds (ridges may not be expressed on external surfaces; Cotton, 2001, p1. 4, fig. 10): C. terranovicus consistently displays a more densely-packed sculpture of fine tubercles and lacks the weak palpebral ridge in large holaspids (Fig. 6.1, 6.3, 6.4, 6.6). In addition, well-preserved specimens demonstrate that the lateral cranidial border of C. terranovicus is reduced to a very narrow strip behind glabellar midlength (Fig. 6.4, 6.6), whereas the lateral border of C. matthewi maintains an even width back to the posterior border furrow (Fig. 5.1, 5.2, 5.6, 5.8; Cotton, 2001, p1. 4, figs. 10, 11). The lateral border furrow of C. terranovicus is broad and maintains an even width, merging with the posterior border furrow (Fig. 6.1, 6.6); lateral and posterior border furrows of C. matthewi do not join (Fig. 5.1, 5.2, 5.6; Cotton, 2001, pl. 4, figs. 10, 11). Dean (in Martin and Dean, 1988) suggested that C. matthewi had a narrower, less tapered glabella than C. terranovicus. However, the material illustrated here (Figs. 5.1, 5.2, 5.5-5.10, 6.1-6.7) demonstrates that both species are variable in glabellar outline, and specimens that are least compacted show minimal differences (compare Figs. 5.6, 5.9 and 6.6). It is possible that C. matthewi and C. terranovicus will prove to represent geographic variation in a single species, and it should also be noted that a poorly known, but apparently very similar, species, C. solvensis (Hicks; Lake, 1940, pt. 61, figs. 14, 15), occurs in the Middle Cambrian of Avalonian Britain.

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Ctenocephalus exsulans (Linnarsson; Westergard, 1950, fig. 9a-d), from the Middle Cambrian Exsulans Limestone of Sweden, has an anteriorly rounded, conical glabella. Other comparisons are difficult because of the compacted nature of the New Brunswick material, but C. exsulans appears to have a lateral border that is reduced to a very narrow (tr.) strip by the submarginal suture, whereas the border is wider in C. matthewi. Ctenocephalus antiquus Thoral (1946, p1. 8, figs. 2, 4, 6; Sdzuy, 1961, p1. 33, figs. 7-9), from the Middle Cambrian of France and Spain appears to differ from all other species by retaining a slender occipital spine in larger holaspids. Like C. matthewi, C. probus (Tchernysheva; Egorova et al., 1982, pi. 1, figs. 9, 10), from the Middle Cambrian of Siberia, has palpebral ridges and sculpture of scattered tubercles. It differs in that the lateral borders are reduced to narrow strips and appear to terminate at the level of the SI glabellar furrow. CTENOCEPHALUS TERRANOVICUS (Resser, 1937) Figure 6.1-6.7 Ctenocephalus terranovicus RESSER, 1937, p. 41, p1. 7, figs. 13, 16, 17. Hartella terranovica (Resser). DEAN in MARTIN AND DEAN, 1988, p. 20, pi. 2, figs. 1, 2, 4-6, 8, 10 (see for synonymy). Diagnosis.-A species of Ctenocephalus that lacks fixigenal ridges and an occipital spine; palpebral ridges absent on internal molds of large holaspids. Glabella subtrapezoidal in outline. Lateral border reduced to narrow strip behind cranidial mid-length; lateral border furrow joins posterior border furrow. Internal molds of cranidia with sculpture of closely spaced tubercles. Material examined.-Twelve cranidia. Occurrence.-Fossil Brook Member, southeastern Newfoundland, collection RBCB 49.5-50.2. Discussion.-Differences between Ctenocephalus terranovicus and C. matthewi, from the Fossil Brook Formation of New Brunswick, were discussed above, and these species are so similar that a description is unnecessary. Most of the criteria used to separate C. matthewi from other members of the genus (discussed above under that species) are equally applicable to C. terranovicus. A small holaspid (Fig. 6.7) differs from co-occurring larger individuals by possession of a stout, subtriangular occipital spine and weak, caecate palpebral ridges. These differences are interpreted as ontogenetic variability, although more material will be need to evaluate this hypothesis fully. Superfamily PARADOXIDOIDEA Hawle and Corda, 1847 Family PARADOXIDIDAE Hawle and Corda, 1847 Discussion.-The most recent evaluation of paradoxidid genera was by Dean and Rushton (1997), who followed concepts laid out by Snajdr (1957, 1958). However, diagnoses provided by Dean and Rushton are essentially descriptive in nature and it is far from clear as to how many of the genera that they recognize are supported by unambiguous synapomorphies. As noted by Geyer and Landing (2001), more than 140 species and subspecies of Paradoxides s.l. have been named, so that a cladistic analysis of the Paradoxididae is well beyond the scope of this paper. Generic groupings used in this paper are essentially those of Dean and Rushton, but we agree with Geyer's (1993, 1998) comments on the need for a major revision of the family. Paradoxides is unique among paradoxidid genera in that the hypostomal suture is non-functional (Snajdr, 1987), and the hypostome is fused to the rostral plate (Snajdr, 1958, pl. 14, figs. 15; Hutchinson, 1962, pl. 22, fig. 3). As several authors have noted (e.g., Snajdr, 1987), the posteriormost thoracic segment is greatly elongated and extends almost straight back and far beyond the posterior end of the pygidium (e.g.. Westergard, 1953, pl. 8, fig. 2; Snajdr, 1958, pl. 15, figs. 1, 4; Hutchinson, 1962, p1. 22, fig. 1). This apparently apomorphic state contrasts with the condition in other paradoxidid genera, including Acadoparadoxides (e.g., Geyer, 1993, pi. 6, pl. 11, fig. 3; Geyer and Landing, 2001, figs. 6.8, 6.10, 7.6; Westergard, 1936, pl. 2, figs. 4, 8-11), Eccaparadoxides (e.g., Fig. 7.1, 10.7, 10.9; Snajdr, 1958, pl. 22, figs. 1, 5, 15), Hydrocephalus (e.g., Snajdr, 1958, pl. 17, fig. 3, pl. 18, fig. 1) and Anabarites (e.g., Dean and Rushton, 1997, fig. 307.2b), in which the posteriormost thoracic segment is considerably shorter. Plutonides (e.g., Hutchinson, 1962, pi. 18, figs. 8, 9) is characterized by very short palpebral lobes centered opposite S2 glabellar furrows, and a glabella that overhangs the anterior border in mature holaspids. Hydrocephalus is also a relatively short-eyed paradoxidid with wide posterior fixed cheeks (e.g., Dean and Rushton, 1997, fig. 304.3a). It possesses a potential apomorphy in the strongly advanced genal spines (e.g., Dean and Ruston, 1997, fig. 304.3b) that were emphasized in Dean and Rushton's (1997, p. 476) diagnosis. Eccaparadoxides and Acadoparadoxides are closely similar taxa with long palpebral lobes that extend from occipital to S4 furrows, and have been variously considered to be separate genera, subgenera or synonyms (see Nikolaisen and Henningsmoen, 1987, p. 67 for discussion). Here, we follow Dean and Rushton (1997) and treat them somewhat arbitrarily as distinct genera. Recent attempts to produce even provisional diagnoses (Snajdr, 1985; Geyer, 1993, 1998; Dean and Rushton, 1997) have met with mixed success, and diagnostic criteria have varied between authors. Geyer's (1998, p. 387; see also Geyer and Landing, 2001) concept of Acadoparadoxides is used here, although some of the characters that he listed are shared with Eccaparadoxides. Criteria suggested by Geyer (1998) as diagnostic of Acadoparadoxides include length of the palpebral lobes, presence of distinct SI and S2 furrows, poorly segmented pygidial axis that occupies more than half of pygidial length and width, and a frontal

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area whose width is similar to cranidial width across the palpebral lobes. Only the latter feature seems to discriminate this genus from Eccaparadoxides on a consistent basis. Species of Eccaparadoxides possess narrow frontal areas that are equal to about 85 percent or less of the cranidial width across the palpebral lobes (e.g., Figs. 8.1, 8.4, 8.5, 9.1, 9.3, 9.8. 10.1, 10.4: see also Dean and Rushton, 1997, fig. 307.1a; Snajdr, 1986, p1. 1, figs. 2, 7, pi. 2, fig. 5). Palpebral lobe length is shared between Acadoparadoxides and Eccaparadoxides, and some species of the latter (e.g., Figs. 8.1, 8.5, 9.1, 9.3) have S3 and S4 furrows that become very shallow in large individuals. Pygidia attributed to Eccaparadoxides are variable, but some (e.g., Figs. 7.1, 8.10, 8.12-8.14) are comparable to those of, for example, Acadoparadoxides briareus Geyer (1993, pl. 10, figs. 1, 3) in the size of the axis. Genus EcCAPARADOXIDES najdr, 1957 Type species*Paradoxides pusillus Barrande, 1846, from the Jince Formation, Czech Republic (by original designation). Remarks.Eccaparadoxides is used somewhat arbitrarily herein to accommodate long-eyed species with relatively narrow frontal areas. ECCAPARADOXIDES ETEMINICUS (Matthew, 1883) Figures 7.1, 7.2, 8.1-8.14, 9.1-9.3 Paradoxides eteminicus MATTHEW, 1883, p. 92, figs. 7-12; HUTCHINSON, 1962, p. 114, pl. 19, figs. 3-8 [only; fig. 9 =Eccaparadoxides acadicus (Matthew)] (see for synonymy). Paradoxides eteminicus Matthew var. suricoides MATTHEW, 1883, p. 97, figs. 4-6. Paradoxides eteminicus Matthew var. breviatus MATTHEW, 1883, p. 99, figs. 1-3. Paradoxides eteminicus Matthew var. malicitus MATTHEW, 1883, p. 101, fig. 13. Paradoxides eteminicus Matthew var. quacoensis MATTHEW, 1883, p. 102, fig. 14, 14a. Paradoxides eteminicus Matthew var. pontificalis MATTHEW, 1883, p. 102, fig. 15, 15a. Paradoxides (Eccaparadoxides) eteminicus (Matthew). DEAN in MARTIN AND DEAN, 1988, p. 18, pl. 1, figs. 2-7, 11, 12, 15. Diagnosis.-A species of Eccaparadoxides with axis that occupies about 75 perecent of pygidial length in large holaspids. S3 and S4 furrows shallow in late meraspids and holaspids. Lectotype (selected herein).-A cranidium (ROM 54082) from Middle Cambrian of New Brunswick (Fig. 8.5). Description.--Cranidium subpentagonal in outline, with length about 85 percent of width across palpebral lobes in uncompacted material; width of frontal area is less than width across palpebral lobes. Convex, bulb-shaped glabella is outlined by finely etched axial and preglabellar furrows, and occupies about 95 percent of cranidial length; expands most rapidly in front of L2 to reach maximum width near S4 furrow. Four pairs of glabellar furrows, with SI and S2 transglabellar on at least exfoliated specimens; SI and S2 well developed, subtransverse to weakly arched backwards; S3 and S4 very shallow and oblique, even in small (?late meraspid) cranidia (Fig. 8.11). Occipital furrow firmly impressed and subtransverse, expanded forward medially and curved gently forward distally. Occipital ring occupies about 15 percent of glabellar length. Anterior border short medially but lengthens abaxially; preglabellar furrow merges with anterior border furrow medially. Palpebral lobe large, arcuate extending from occipital furrow to S4; palpebral area of fixigenae gently convex on uncompacted specimens and wide, with maximum width equal to about half of glabellar width at L2 lobe. Anterior branches of facial suture diverge sharply forward before swinging inward along strongly curved anterior cranidial margin; posterior branches short, straight, and divergent. Posterior border furrow well defined and isolates short, convex posterior border from remainder of fixigenae. On testate specimens (Fig. 9.1-9.3), anterior half of glabella and anterior border carry sculpture of fine terrace ridges. Posterior half of glabella and palpebral area of fixigenae are finely granulose; granules augmented by fine caecal network on fixigenae, and oblique ridge extends forward from anterior end of palpebral lobe to anterior border. Librigena with long, stout genal spine equal to about half of thoracic length. Lateral borders and doublure broad, border furrows shallow. Internal molds smooth, except for terrace ridges on doublure. Hypostome not fused with rostral plate, loaf-shaped in outline with transverse posterior margin and broadly rounded anterior margin. Median body differentiated into two lobes by median furrow that is faint medially but deepens at maculae. Anterior lobe large, convex; posterior lobe crescentic, and weakly convex. Anterior wings short and triangular. Lateral and posterior borders narrow, separated from middle body by moderate deep border furrow; junctions of lateral and posterior borders extended into long, weakly incurved spines. Thorax with 17 segments ending in long pleural spines that occupy more than half of pleural lobe width. Axis evenly and slightly tapered backwards occupying about 33 to 40 percent of thoracic width. Axial ring with short, forwardly curved articulating half-ring and subtranverse articulating furrow. Inner pleural region more or less transverse and narrower (tr.) than axis; outer pleural region and spine curved gently backward. Pleural furrows diagonal, deepest medially and narrowing towards both axis and spine. Pygidium subhexagonal in outline, with straight or concave posterior margin; some specimens (Fig. 8.7, 8.8) have pointed posterolateral tips. Larger pygidia (Fig. 8.10) are more rounded in outline than smaller specimens (Fig. 8.12, 8.13). Convex axis is longer than wide and tapers

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backwards for about four-fifths of pygidial length. Up to two axial rings and long terminal piece evident and best defined on exfoliated surfaces; at least anteriormost axial ring well-defined and subtransverse. First axial ring with narrow articulating half-ring and transverse articulating furrow at anterior, and depressed crescentic region at posterior. Pleural field largely effaced, with one or two oblique, poorly defined pleural furrows. Border not differentiated from effaced pleural field. Doublure very wide with sculpture of concentric terrace ridges. External surface covered with closely-spaced, small, pointed or rounded granules (Fig. 8.7, 8.14), whereas internal molds are smooth (Fig. 8.13). Material examined.-Two complete exoskeletons, one hundred and thirteen cranidia and twenty-two pygidia from field collections; twelve cranidia and one librigena from Matthew's types; eight cranidia from NYSM collections. Occurrence.--Fossil Brook Member, Chamberlain's Brook Formation, New Brunswick (collection FoB-FB 0.2, 2.5; 11314.85, 15.5; GoS 2.3) and Newfoundland (collection RBCB 49.550.2). Matthew's types were collected at Saint John, probably at Seely Street; NYSM material is from Seely Street. Discussion.-A nearly complete exoskeleton from Newfoundland (Fig. 7.1) provides the first unequivocal association of sclerites of Eccaparadoxides eteminicus. Pygidia are subhexagonal in outline with a relatively long axis. Among the pygidia assigned by Hutchinson (1962, pl. 19, fig. 9) to this species is a specimen with a short axis, and this may belong to E. acadicus (see following discussion of E. acadicus). In collections from Newfoundland, pygidia of E. eteminicus (Fig. 8.10, 8.12-8.14) are variable and have transverse or gently embayed posterior margins, with rounded or sharply pointed (Fig. 8.7, 8.8) postero-lateral margins; external surfaces have granulose sculpture (Fig. 8.7, 8.8, 8.14) and internal molds are smooth (Fig. 8.13). Matthew (1883) established several varieties of E. eteminicus (e.g., see Walcott, 1884, pl. 3, fig. 1, lc-d) for cranidia from New Brunswick. They fall within the range of variability of material in our collections (in part due to compaction and/or tectonic shear), and all are included within E. eteminicus. In possessing weak S3 and S4 furrows, the cranidium of Eccaparadoxides eteminicus is similar to E. asturianus Sdzuy (1967, p1. 2, figs. 1-5) from the Middle Cambrian of Spain, differing in having a shorter anterior border. In addition, the pygidium of E. asturianus has a shorter axis and more convergent lateral margins. producing a more strongly hexagonal outline (Sduzy, 1967, pl. 21 figs. 4, 5). There appears to be little to separate cranidia of a second Spanish species, E. sdzuyi Linan (e.g., Lin et al., 1995, pl. 1, figs. 2-6) from those of E. eteminicus. Like F. eteminicus, pygidia of E. sd Eccaparadoxides acadicus (Matthew- Fig. 10.1, 10.3, 10.4, 10.6), E. rohanovicus Snajdr (1985, pl. 1, figs. 1-7, pl. 2, figs. 4-6) and E.? insularis (Westergard, 1936, pl. 7, figs. 1-9) are differentiated readily from E. eteminicus by the presence of firmly impressed S3 and S4 glabellar furrows, whereas E. lamellatus (Hartt; Fig. 10.9-10.11) possesses a distinctive sculpture of conspicuous, raised ridges on the anterior glabellar lobe, with coarse granules on the remainder of the glabella. The type species of Eccaparadoxides, E. pusillus (Barrande) has well-defined S3 and S4 furrows on small cranidia (Snajdr, 1958, p1. 21, fig. 4; 1985, pl. 2, fig. 1) although, like E. eteminicus, they are weak at larger sizes (Snajdr, 1985, pi. 2, figs. 2, 3). The hypostome of E. pusillus has shorter, divergent spines on posterior corners (Snajdr, 1958, p1. 21, figs. 8, 16-18); a similar hypostome to that of E. eteminicus was attributed to Acadoparadoxides sirokyi Snajdr by Snajdr (1987, pl. 2, fig. 7). Pygidia have a more pronounced hexagonal outline and a much shorter axis that occupies less than half of pygidial length (Snajdr, 1958, p1. 22, figs. 9-15). Paradoxides (Eccaparadoxides) remiss Dean possesses narrow inter-occular fixigenae, so that width across the palpebral lobes is slightly less than maximum width of the frontal area (Dean, 1982, fig. 27). As such, it does not conform to Eccaparadoxides as used here, and cranidial proportions are similar to those of Acadoparadoxides. However, pygidia attributed to this species by Dean (1982, figs. 23, 32, 34) are very long and narrow with short axes, and differ from those of species currently included in Acadoparadoxides (e.g., Snajdr, 1958, pl. 22, figs. 9-15; Geyer, 1993, pi. 10, figs. 1, 3; Westergard, 1936, pl. 3, figs. 5-16). The assignment of this species is uncertain, although it appears to be part a group of peri-Gondwanan species, variously placed in Paradoxides s.1. or Eccaparadoxides (e.g., Dean, 1982, fig. 86; Sdzuy, 1961, figs. 26, 27, p1. 19, figs, 14-18; pi. 21, figs. 3-9), that share similar pygidial morphology. ECCAPARADOXIDES cf. ETEMINICUS (Matthew, 1883) Figure 9.4-9.11 Material examined.-Seven cranidia and three pygidia. Occurrence.-Fossil Brook Member, Chamberlain's Brook Formation, New Brunswick, collection HBW-33.2-33.85. Discussion.-A few cranidia (Fig. 9.5, 9.8-9.11) preserved in full relief in the basal limestone bed at Hanford Brook resemble uncompacted specimens of E. eteminicus (Fig. 9.1-9.3) in possessing weak S3 and S4 furrows. They differ in having less strongly curved anterior cranidial margins and glabellae that are more expanded anteriorly. Associated pygidia (Fig. 9.4, 9.6, 9.7) are subquadrate in outline and are quite distinct from those of E. eteminicus (Fig. 8.7, 8.8, 8.10, 8.12-8.14). In addition, the axis is relatively wider and unsegmented, whereas even small pygidia of E. eteminicus (Fig. 8.8) display at least one well-defined axial ring. The Hanford Brook material probably represent a new species but are best placed in open nomenclature until more specimens become available. ECCAPARADOXIDES ACADICUS (Matthew, 1883)

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Figure 10.1-10.8 Paradixides acadicus MATTHEW, 1883, p. 103-104, figs. 16-18; WAL COTT, 1884, p. 25-27, pl. 3. 3a: MATTHEW, 1884, p. 99, figs. 1, 2. Paradoxides eteminicus Matthew. HUTCHINSON, 1962. p. 114, pl. 19, fig. 9 [only]. Diagnosis.-A species of Eccaparadoxides with deeply incised S3 and S4 glabellar furrows and large median node on occipital ring. Anterior cranidial border narrows medially. Cranidium has coarsely granulose sculpture. Lectotype (selected herein).-A cranidium (ROM 54095) from the Fossil Brook Member, Chamberlain's Brook Formation, New Brunswick (Fig. 10.4). Description.-Cranidium subpentagonal in outline; cranidial width is greatest between palpebral lobes. Convex, anteriorly rounded glabella expands forward to reach maximum width at S4 glabellar furrow; occupies about 92 percent of cranidal length and about 40 percent of cranidial width across palpebral lobes. Four pairs of firmly impressed glabellar furrows; SI is transglabellar although shallow medially, and bowed gently backwards; subtransverse S2 also transglabellar and shallowing medially; S3 discontinuous, curving gently forward from axial furrow: S4 discontinuous and transverse. Occipital furrow deep and transverse near axial furrow, becoming very shallow and forwardly convex medially. Occipital ring with large median node. Anterior border widest at anterior corners of cranidium and narrowing somewhat medially; preglabellar furrow merges with anterior border furrow medially. Long, arcuate palpebral lobe extends from occipital furrow to S4 furrow; palpebral furrow broad, shallow but clearly defined. Anterior branches of facial suture initially divergent before curving inward along anterior cranidal margin; posterior branches short, straight and gently divergent. Posterior border short and curved gently backwards; border furrow firmly impressed. Fixigenae and glabella with coarsely granulose sculpture. Librigena with long, stout genal spine. Lateral border and doublure broad, equal to slightly more than half of librigenal width. Doublure carries conspicuous terrace ridges. Thorax (Fig. 10.7) has at least twelve segments that terminate in backwardly curved spines, with the amount of curvature increasing progressively backwards. Narrow axis tapers backwards, accounting for only about twenty percent of thoracic width towards anterior; axial furrows shallow but clearly defined. Axial ring includes short, subtransverse articulating half-ring and firmly impressed articulating furrow. Each pleura carries well incised, oblique pleural furrow that occupies slightly less than 30 percent of pleural width. Pleurae decrease in length towards rear, and two posteriormost pleurae curve almost directly backwards to terminate a short distance beyond the posterior tips of the pygidia. Pygidium elongate, with maximum width about 70 percent of length, and subhexagonal in outline; posterior margin is embayed and postero-lateral corners carry short triangular spines. Axis short, equal to about 45 percent of pygidial length, subtriangular in outline with rounded posterior termination. Short, transversely semielliptical articulating half-ring separated from remainder of axis by deep articulating furrow. One axial ring differentiated and has short, narrow sub-semielliptical depressed area at posterior; transverse axial ring furrow shallow but clearly defined. Second, poorly defined axial ring and shallow, incomplete ring furrow may be evident on some specimens. Doublure long, extending to axis and carries terrace ridges. Material examined.-Four cranidia from field collections plus five cranidia, one pygidium, two librigena, and a thoracopygon. Occurrence.-Fossil Brook Member, Chamberlain's Brook Formation, New Brunswick and Newfoundland (collection RBCB 49.5-50.2). Matthew's types are from New Brunswick (Seely Street, Saint John), and the thoracopygon attributed to E. acadicus herein (Fig. 10.7) is from St. Martins, New Brunswick. Discussion.-The pygidium in the type lot of Eccaparado-vides acadicus (Fig. 10.2) is incomplete, but has outwardly curved lateral margins and a short axis that appears to have occupied less than half of pygidial length. It is very similar to pygidia from Fossil Brook (Fig. 10.8) which are, in turn, closely comparable to a pygidium and associated thorax (Fig. 10.7) among the referred specimens of E. lamellatus (Hartt) in the collections of the Royal Ontario Museum. The latter specimen differs in several features from a second exoskeleton that may be attributed confidently to E. lamellatus (Fig. 10.9) and is tentatively transferred to E. acadicus (see following discussion of E. lamellatus). Finally, a pygidium identified as E. eteminicus by Hutchinson (1962, pi. 14, fig. 9) closely resembles the referred specimen (Fig. 10.7), and it is also assigned to E. acadicus. The most similar species to E. acadicus is E. rohanovicus Snajdr, 1986 (PI. 1, figs. 1-7, pi. 2, figs. 4-6) from the Middle Cambrian of Bohemia. Like E. acadicus, E. rohanovicus has welldefined S3 and S4 furrows throughout holaspid ontogeny, and differs in having more strongly curved palpebral lobes, a longer anterior cranidial border that is not reduced in width medially, and a more finely granulose sculpture. The pygidium of E. rohanovicus (e.g., Snajdr, 1986, fig. lb) is relatively wider than those assigned to E. acadicus (Fig. 10.2, 10.7, 10.8) and has a more deeply embayed posterior margin with longer marginal spines. Eccaparadoxides pusillus (Barrande), also from Bohemia, has pooly defined S3 and S4 lateral furrows on larger individuals (Snajdr, 1986, pl. 1, figs. 1-3). The pygidium of E. pusillus (e.g., Snajdr, 1986, fig. la) is similar to that of E. acadicus but has a subtransverse posterior margin with longer marginal spines. ECCAPARADOXIDES LAMELLATUS (Hartt in Dawson, 1868) Figure 10.9-10.11

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Paradoxides lamellatus HARTT in DAWSON, 1868, p. 656-657; WALCOTT, 1884, p. 25, pi. 3, fig. 2, 2a (see for synonymy); SHIMER AND SHROCK, 1944, p. 615, pl. 254, figs. 14, 15; pl. 276, fig. 19; HUTCHINSON, 1962, p. 113, pl. 18, figs. 1-3 (see for synonymy). Diagnosis.-A species of Eccaparadoxides having raised ridges and irregular tubercles on L4 and frontal glabellar lobes; remainder of glabella with coarsely granulose sculpture. Material examined.-Two cranidia from field collections and three, variously complete exokeletons from the Matthew Collection, ROM. Occurrence.-Fossil Brook Member, Chamberlain's Brook Formation, New Brunswick, collection FoB-FB 0.2. Matthew's specimens are from St. Martins, New Brunswick. Discussion.-The cranidia in our collections are incomplete but are clearly conspecific with Hart's types, as figured by Walcott (1884, pl. 3, fig. 2, 2a). The raised ridges and irregular tubercles on the L4 and frontal glabellar lobes are not true terrace ridges because they are expressed on internal molds. In this respect, they resemble the striate ridges and irregular tubercles on some species of the Late Cambrian dikelocephalacean Ptychaspis (e.g., Westrop, 1986, pl. 8, figs. 1, 2, 9). A nearly complete exoskeleton (Fig. 10.9) is a cast from an external mold and compares favorably with the individual illustrated by Matthew (1890, pl. 11, fig. 9). It is not known whether this is the actual figured specimen or simply the counterpart but, in any event, it is the only complete exoskeleton in the collections at the Royal Ontario Museum. The pleurae are relatively narrow (tr.), and the posteriormost pair do not extend back beyond the rear of the pygidium. The pygidium is incomplete posteriorly, but the unsegmented axis appears to have occupied at least half of its length. A pygidium attributed to E. lamellatus by Hutchinson (1962, pl. 18 fig. 3; this specimen is from a different locality than the cranidia figured by Hutchison, 1962, pi. 18, figs. 1, 2) has a shorter axis but also appears to be unfurrowed. A smaller, incomplete thoracopygon (Fig. 10.7) was also present among the specimens identified as E. lamellatus at the Royal Ontario Museum, and this has much longer pleurae, with the three posteriormost pairs extending beyond the rear of the pygidium. The pygidium is complete and has a short axis that has a well-defined axial ring. As noted above, it closely resembles the syntype pygidium of E. acadicus, and is tentatively assigned to that species. ACKNOWLEDGMENTS Fieldwork was supported by National Science Foundation grants EAR94-12577 and EAR98-05177 to E. Landing, and Natural Sciences and Engineering Research Council of Canada Research Grant 41197 to S. R. Westrop. D. H. Kim was supported by Korean Research Foundation Grant KRF-99-D042 and the BK 21 Project. D. M. Rudkin and J. Waddington arranged for the loan of type and referred specimens from the Matthew Collection at the Royal Ontario Museum, and P Krohn attempted to locate Hartt's types in collections of the Paleontological Research Institute. A. R. Palmer and A. W. A. Rushton provided helpful reviews of the manuscript, and Palmer generously provided copies of pertinent Russian literature. REFERENCES ANGELIN, N. P 1854. Paleontologia Scandinavica. Pars 1. Crustacea Formationis Transitionis. Fasc. II. Academiae Regiae Scientarium Suecanae: I-IX + 21-92, T 0. Weigel, Leipzig. ANGEt.IN, N. P 1878. Paleontologia Scandinavica. Appendix, 93-96. Holmiae, Stockholm. BABCOCK, L. E. 1994. Systematics and phylogenetics of polymeroid trilobites from the Henson Gletscher and Kap Stanton formations (Middle Cambrian), North Greenland. Gronlands Geologiske Undersoglese Bulletin, 169:79-127. BARRANDE, J. 1846. Nouveaux Trilobites. Supplement a la notice preliminaire sur le systeme silurien et les Trilobites de Boheme. Prague and Leipzig, 97 p. COTTON, T. J. 2001. The phylogeny and systematics of blind Cambrian ptychoparioid trilobites. Palaeontology, 44:167-207. DAWSON, J. W. 1868. Acadian Geology: The Geological Structure, Organic Remains and Mineral Resources of Nova Scotia, New Brunswick, and Prince Edward Island. London. DEAN, W. T. 1982. Middle Cambrian trilobites from the Sosink Formation, Derik-Mardin district, south-eastern Turkey. Bulletin of the British Museum (Natural History), Geology Series, 36:1-41. DEAN, W. T., AND A. W. A. RusHTON. 1997. Superfamily Paradoxidoidea Hawle & Corda, 1847, p. 470-481. In H. B. Whittington et al., Treatise on Invertebrate Paleontology, Pt. 0, Arthropoda. Trilobita, Revised, Volume 1. The Geological Society of America and The University of Kansas Press. Lawrence. EGOROVA, L. I., Y. Y SHABANOV, T V. PEGEL, V E. SAVITSKY, S. S. SUCHOV, AND N. E. TCHERNYSHEVA. 1982. Maya Stage of type locality (Middle Cambrian of Siberian platform). Academy of Sciences of the USSR, Ministry of Geology, Interdepartmental Stratigraphic Committee, Transactions, 8, 146 p. (In Russian) EGOROVA, L. I., YU. YA. SHABANOV, A. Yu. ROZANOV, V. E. STAVITSKY, N. E. TCHERNYSHEVA, AND B. B. SHISHKIN. 1976. Elanka and Kuonamka facies stratotypes of the lower boundary of the Middle Cambrian in Siberia. Nedra, Moscow. 168 p. (in Russian)

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MATTHEW, G. F 1883. Illustrations of the fauna of the St. John Group. No. 1. The Paradoxides. Transactions of the Royal Society of Canada, 2(4):271-279. MATTHEW, G. E 1885 (1884). Illustrations of the fauna of the St. John Group continued: No. 2: On the Conocoryphaea, with further remarks on Paradoxides. Transactions of the Royal Society of Canada, 3(4): 99-124. MATTHEW, G. E 1889. On Cambrian organisms in Acadia. Transactions of the Royal Society of Canada, 7(4):135-162. MATTHEW, G. F 1890. Illustrations of the fauna of the St. John Group, No. V. Transactions of the Royal Society of Canada, 4, 123-166. MCCARTNEY, W. D. 1967. Whitbourne Map-Area, Newfoundland. Geo logical Survey of Canada, Memoir 341, 135 p. MILLER, S. A. 1889. North American Geology and Paleontology. Cincinnati, Ohio, 664 p. MYROW, P. M., AND E. LANDING. 1993. Mixed siliciclastic-carbonate deposition in an Early Cambrian stratified basin, Chapel Island Formation, southeastern Newfoundland. Journal of Sedimentology Petrology, 62:455-473. NIKOLAISEN, E, AND G. HENNINGSMOEN, 1987. Lower and Middle Cambrian trilobites from the Digermul Peninsula, Finmark, northern Norway. Norges geologiske undersokelse Bulletin, 419:55-95. POULSEN, C. 1959. Family Conocoryphidae, p. 0242-0244. In R. C. Moore (ed.), Treatise on Invertebrate Paleontology, Pt. 0, Arthropoda 1. Geological Society of America and University of Kansas Press, Boulder and Lawrence. RESSER, C. E. 1936. Second contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 95, No. 4, 29 p. RESSER, C. E. 1937. New species of Cambrian trilobites of the family Conocoryphidae. Journal of Paleontology, 11:39-42. SALTER, J. W. 1864. A monograph of the British trilobites from the Cambrian, Silurian and Devonian formations. Palaeontographical Society Monograph, 80 p. SALTER, J. W., AND H. HICKS. 1869. On some fossils from the "Menevian Group". Quarterly Journal of the Geological Society of London, 25: 51-57. SDZUY, K. 1958. Neue Trilobiten aus dem Mittelkambrium von Spanien. Senkenbergiana Lethaia, 39:235-253. SDZUY, K. 1961. Das Kambrium Spaniens. Teil 11: Trilobiten, 1. Akademie der Wisseschaften and der Literatur in Mainz, Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse, 7:499-594. SDZUY, K. 1967. Trilobites del Cambrico Medio de Asturias. Trabajos de Geologia, 1:77-133. SDZUY, K., E. LiNAN, AND R. GOZALO. 1999. The Leonian Stage (early Middle Cambrian): a unit for Cambrian correlation in the Mediterranean subprovince. Geological Magazine, 136:39-48. SHIMER, H. W., AND R. R. SHROCK. 1944. Index Fossils of North America. The Technology Press, Massachusetts Institute of Technology, Cambridge, Massachusetts, 837 p. SKEHAN, J. W., D. P. MURRAY, A. R. PALMER, A. T. SMITH, AND E. S. BELT. 1978. Significance of fossiliferous Middle Cambrian rocks of Rhode Island to the history of the Avalonian microcontinent. Geology, 6:694-698. SNAJDR, M. 1957. 0 novych trilobitech z ceskeho kambria. Vestnik Ustredniho fistavu Geologicky, 32(4):235-244. (In Czech, with English summary) SNAJDR, M. 1958. Trilobiti aeskrho stredniho kambria. Rozpravy Ustredmho dstava geologickeho, 24, 280 p. (In Czech, with English summary) SNAJDR, M. 1986. Two new paradoxidid trilobites from the Jince For mation (Middle Cambrian, Czechoslovakia). VGstnik Ustrfdniho astavu geologickych, 61(3):169-174. SNAJDR, M. 1987. The genera Paradoxides Brongniart and Hydrocephalus Barrande (Trilobita). Vestnik Ustredniho ustavu geologickeho, 62(2):97-104. THORAL, M. 1946. Conocoryphidae Languedociens. Annales de l'Universit' de Lyon, Troisieme Serie, Section C, Sciences Naturelles, 4:5-74. WALCOTT, C. D. 1884. On the Cambrian faunas of North America. United States Geological Survey, Bulletin 10, pp. 289-355. WESTERGARD, A. H. 1936. Paradoxides oelandicus Beds of ()land with the account of a diamond boring through the Cambrian at Mrssberga. Sveriges Geologiska Undersokning, Series C, 394:1-66. WESTERGARD, A. H. 1950. Non-agnostidean trilobites of the Middle

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Cambrian of Sweden. II. Sveriges Geologiska Undersokning, Series C, 43, 57 p. WESTERGARD, A. H. 1953. Non-agnostidean trilobites of the Middle Cambrian of Sweden. 111. Sveriges Geologiska Undersokning, Series C, 46, 59 p. WESTROP, S. R. 1986. Trilobites of the Upper Cambrian Sunwaptan Stage, southern Canadian Rocky Mountains, Alberta. Palaeontographica Canadiana, 3, 175 p. WESTROP, S. R., AND E. LANDING. 2000. Lower Cambrian (Branchian) trilobites and biostratigraphy of the Hanford Brook Formation, southern New Brunswick. Journal of Paleontology, 74:858-878. WHITEAVES, J. E 1878. On some Primordial fossils from southeastern Newfoundland. American Journal of Science, 3, Series, 16:224-226, ZHANG, W. T, AND P A. JELL. 1987. Cambrian Trilobites of North China. Science Press, Beijing, 459 p. ACCEPTED 3 OCTOBER 2001 DONG HEE KIM,1 STEPHEN R. WESTROP AND ED LANDING2 Oklahoma Museum of Natural History and School of Geology and Geophysics, University of Oklahoma, Norman 73072, and 2Center for Stratigraphy and Paleontology, New York State Museum, The State Education Department, Albany 12230 1 Present address: School of Earth and Environmental Sciences, College of Natural Sciences, Seoul National University, Seoul 151-742, Korea. Copyright Paleontological Society Sep 2002 Provided by ProQuest Information and Learning Company. All rights Reserved

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