THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS

May 16, 1978 Paper 90

REVISION OF A LATE MIDDLE CAMBRIAN TRILOBITE FAUNULE FROM NORTHWESTERN QUEENSLAND'

P. A. JELL and R. A. ROBISON University of Queensland, St. Lucia, Queensland, and University of Kansas, Lawrence

ABSTRACT Three species, each belonging to a different genus, are shown to be represented by previously figured types of Euagnostus opimus Whitehouse, 1936, which is the nominate species of the commonly cited Euagnostus opimus Zone in Australia. The holotype of E. opimus is here reassigned to Peronopsis Corda, 1847, and Euagnostus Whitehouse, 1936, is thereby suppressed as a subjective junior synonym of Peronopsis. This action necessitates a change in biostratigraphic terminology from Euagnostus opimus Zone to Peronopsis opimus Zone. Because of the taxonomic and biostratigraphic importance of these conclusions, all trilobites from the type collection of E. opimus are described or redescribed and evaluated. These include the miorneroids Baltagnostus australis n. sp., Doryagnostus deltoïdes n. sp., Hypagnostus? sp., Opsidiscus micros pinus Jell, and Peronopsis opimus (Whitehouse) as well as the polymeroids Chondranomocare confertum (Whitehouse), Penarosa retif era Opik, Prodamesella biserrata n. sp., and Sudanomocarina changi n. gen. and n. sp. Also, type specimens of Baltagnostus damesi (Resser & Endo) from Manchuria are redescribed and refigured for comparison with B. australis, and a lectotype for B. damesi is designated.

INTRODUCTION Euagnostus has been a poorly understood subjective junior synonym of Peronopsis. Figured Middle Cambrian trilobite genus. Its type species, paratypes of E. opimus include a pygidium of a E. opimus, was described in 1936 from north- new species of Baltagnostus Lochman in Lochman western Queensland by the late F. W. White- and Duncan, 1944, and a cephalon of a new house. Recent examination of the type collection species of Doryagnostus Kobayashi, 1939. Revised of E. opimus has led us to the conclusion that the taxonomic assignment of these specimens has ad- holotype and two of Whitehouse's figured para- ditional significance because Opik (1970) has types of E. opimus actually represent three spe- defined an Euagnostus opimus Zone in Australia cies, each belonging to a different genus. We that has been cited in several subsequent publica- have also concluded that the holotype of E. opimus tions (e.g., Hill, Playford, & Woods, 1971; Jell, should be reassigned to Peronopsis Corda (in 1975; Shergold & others, 1976). Because of the Hawle & Corda, 1847), leaving Euagnostus a taxonomic and biostratigraphic importance of

1 Manuscript received June 29, 1977. Whitehouse's material, all trilobites from his

2 The University of Kansas Paleontological Contributions—Paper 90 original collection are described or redescribed field (1959, p. 0117-0126); however, notations and evaluated in this paper. for glabellar segmentation of the polymeroids Whitehouse's collection from the Currant Bush follow Richter and Richter (1940) and some Limestone, which was made in 1932, is housed agnostoid terminology is from Robison (1964). in the Department of Geology and Mineralogy, As used for polymeroids, the term "glabella" in- University of Queensland, and is labeled with cludes the occipital ring. the locality number UQL256. Figured specimens Acknoudedgments.—W. T. Chang of the are entered on the Register of Fossils of the same Nanking Institute of Geology and Palaeontology institution and are identified by numbers with provided photographs of Prodamesella con vexa the prefix UQF. For comparative purposes, a Chang as well as useful correspondence with P. few specimens of Baltagnostus damesi (Resser & A. Jell concerning this faunule. F. J. Collier ar- Endo) are figured from the collections of the U.S. ranged for the loan of specimens from the U.S. Museum of Natural History, Washington, D.C., Museum of Natural History, and A. R. Palmer and these are designated by numbers with the and A. J. Rowell critically reviewed the manu- prefix USNM. script. Research by R. A. Robison was supported Systematic descriptions of the agnostoid trilo- by National Science Foundation grants GA-43723 bites have been prepared by R. A. Robison and and EAR76-10953, and the Wallace E. Pratt Re- those for an eodiscoid and the polymeroids are search Fund, which was provided by Exxon by P. A. Jell. Morphological terms are mostly U.S.A. Foundation and administered by the Uni- those defined by Harrington, Moore, and Stubble- versity of Kansas.

LOCALITY

The exact location of UQL256 is a subject of Whitehouse's field notes in the Fryer Library some confusion (Fig. 1). In the description of of the University of Queensland clear away some the species, Whitehouse (1936, p. 87) gave as of the confusion. His entry for January 14, 1938, the locality for Euagnostus opimus "the Anomo- explains that since he was last in the area, the care Stage, 52 miles from Camooweal on the Thorntonia homestead had been moved from its road from Camooweal to Thorntonia Station," original location on the main Thornton River and in the section of the same paper titled "Notes to a place on the West Thornton River about on the Faunal Stages" he (p. 76) mentioned that 10 miles upstream from its junction with the the Anomocare Stage "at the road crossing of Thornton. Also, the main Camooweal to Burke- Harris Creek north-east of Camooweal" contained town road had been realigned so as still to pass Anomocare, Euagnostus, Phalacroma, and brachi- by the homestead. These two roads and home- opods. This is basically the fauna of UQL256, stead sites are marked on the Camooweal hut whether or not the type locality of the stage is 1 :253,440 and 1:250,000 geological maps (the old geographically separate from that of Euagnostus road is dashed on these maps). opimus was not indicated. In 1939 Whitehouse On January 15, 1938, Whitehouse wrote that (p. 224) gave as the locality for Anon2ocare he "ran onto Harris Creek (locally called the con fertum "the Anomocare Stage, five miles east Douglas) and followed it down to Undilla." He of Harris Creek on the old main road from also mentioned "Magenta Creek (locally known as Camooweal to Burketown, via Thorntonia Sta- Harris)." A search of the state government maps tion (52 miles from Camooweal)." In the same of the area has shown that these local usages sub- paper (p. 270) he listed the coordinates of the sequently received official recognition. The termi- "road crossing of Harris Creek (Camooweal to nology used by Whitehouse in 1936 and 1939 was Old Thorntonia)" as 138°43' E.; 19°26' S. In used as late as 1951 on a map of the state's stock 1945 (p. 121) he mentioned that Dorypyge tenella routes. However, on the 1960 geological map came from beds "about four miles east of Douglas and the 1968 official parish map, the name of Creek on the old Burketown road. (This locality Harris Creek (with Undilla homestead adjacent) is about three-quarters of a mile west of the type is changed to Douglas Creek and the name of the locality for Anomocare confertum.)" more easterly Magenta Creek is changed to Harris

Jell & Robison—Revision of a Trilobite Faunule 3

138°40 E 138°50' E

Old ..\Thorntonia 19°25' 19°25' S

19°35' S — 19°35' S SCALE \-\ 0 Miles 5 i

0 Kilometers 8 (

138°40' E 138°50' E FIG. I. Index map of the Thorntonia area, northwestern Queensland.

Creek. Clearly, Whitehouse used the newer east of Bureau of Mineral Resources (BMR) lo- terminology in his 1945 paper. cality 28 marked on the Camooweal Geological The locality 52 miles from Camooweal is Map (same as M28 on Fig. 1), and the sequence about five miles east of Douglas Creek (formerly from the Ptychagnostus atavus to Ptychagnostus Harris Creek). Whitehouse's field notes for a punctuosus zones is known in several northeast later visit (1939) reveal that the position of the to southwest sections across the Currant Bush road crossing quoted for the Anomocare Stage Limestone from BMR locality 28 to as far as five in 1936 is in error. Coordinates quoted refer to miles south of the present Thorntonia homestead. modern Harris Creek (formerly Magenta) on One new species, Sudanomocarina changi, is the old Camooweal to Burketown road, an area not well represented in Whitehouse's collection of almost unfossiliferous and older limestone, of (UQL256), and supplementary material from a lithology quite different (sandy, dolomitic, al- UQL463 has been used to complete the descrip- most massive) from that of the Anomocare Stage. tion. UQL463 is on top of a hill 5.6 km (3.5 mi) One can only assume that the newly noticed south of Thorntonia homestead on the left bank (1937-38 field notebook) changes in geographic of the West Thornton River and has been located names contributed to Whitehouse's error. and recollected by one of us (P.A.J.). It includes Locality UQL256 is, therefore, reasonably well the same fauna as UQL256 but has some addi- identified and a similar fauna has been collected tional species. To complete the revision of ma- by one of us (P.A.J.) from that general area, but terial referred to Euagnostus by Whitehouse the precise location of Whitehouse's collecting (1939, p. 260) a pygidium, UQF69639, from a site is almost impossible to determine because the locality near the Camooweal to Burketown road limestone strata lie almost flat and consequently crossing of V Creek is also described. are widespread. (Indeed, Whitehouse's 1939 field On the Camooweal Geological Map, localities notes mention stopping at several places that he UQL256 and 463 are in the outcrop area of the took to be the type locality, but which turned out Currant Bush Limestone. Locality UQL463 has to be unfossiliferous or to contain a different also been mapped as such by deKeyser and Cook faunule.) The exact location is somewhere just (1972, p. 27).

4 The University of Kansas Paleontological Contributions—Paper 90

CONTENT, AGE, AND DISTRIBUTION OF FAUNULE

The following trilobite taxa have been identi- Middle Cambrian Hypagnostus parvifrons Zone fied from collection UQL256: of Scandinavia. In the absence of H. parvifrons in Australia, he assigned the equivalent interval Miomeroids to the Euagnostus opimus Zone. Because the nominate species of that zone is here shown to Peronopsis opimus (Whitehouse, 1936) represent three separate species among its pre- Doryagnostus deltoides Robison, n. sp. viously figured types, and because Euagnostus is Baltagnostus australis Robison, n. sp. here suppressed as a subjective junior synonym of Hypagnostus? sp. Peronopsis, the name of the biostratigraphic unit Jell, 1975 Opsidiscus microspinus is here changed to the Peronopsis opimus Zone. Chondranomocare and Sudanomocarina arc Polymeroids present in the upper half of the Amga Stage of Chondranomocare con fertum (Whitehouse, 1939) Siberia in association with Ptychagnostus gibbus Sudanomocarina changi Jell, n. gen. and n. sp. so that the occurrence in UQL256 extends their Prodamesella biserrata Jell, n. sp. known ranges by two agnostoid zones. In Man- Penarosa retif era 6pik, 1970 churia these genera occur in the Mapania Zone. gen. and sp. undet. (dorypygid? cranidium) Prodamesella occurs in the same zone but at dif- ferent localities in China, and apparently is not The fauna, with some additions and deletions, associated with either Chondranomocare or Su- is well known locally from several sections along danomocarina. According to Chang (1957) and the northeastern margin of the Undilla basin, Lu (1960), the Mapania Zone (or lower part of where it overlies a Ptychagnostus atavus fauna the Am photon Zone) may correlate with the and underlies a fauna containing Ptychagnostus Ptychagnostus gibbus Zone, and therefore, the punctuosus. In considering these relationships, Chinese faunas of the Mapania Zone may also Opik (1970, p. 4) recognized that the fauna rep- be slightly older than the Australian faunulc resented the same period of time as the late containing Prodamesella and Sudan omocarina.

SYSTEMATIC DESCRIPTIONS Agnostoid Trilobites median node weak or effaced; posterior end con- [by R. A. Robison I stricted by slight indentations to accommodate inward displacement of simple basal lobes, com- Because of the current inadequacy of agnostoid monly with tiny median ridge above cephalic supergeneric taxa are not classification, names of recess. Genae smooth. Preglabellar median fur- genera are arranged used in this paper, and row incomplete; deepest near glabella, shallower alphabetically. toward anterior, and usually disappears before Genus BALTAGNOSTUS Lochman reaching border furrow. Lateral sides of acrolobes almost parallel or slightly tapered toward anterior. in LOCHMAN & DUNCAN, 1944, p. Baltagnostus LOCHMAN Border furrow wide and border narrow. 138; PALMER, 1955, p. 718; HOWELL, 1959, p. 0184; ROBISON, 1964, p. 525-526; 0Pix, 1967, p. 77. Pygidium subquadrate to transversely subrec- Axial furrow well defined. Axis rela- Type species.—Proagnostus? centerensis RES- tangular. second seg- SER, 1938, p. 48, by original designation. tively broad, slightly constricted at ment, and may or may not extend to border Diagnosis.—Small agnostoids; length of com- furrow; transaxial furrows usually effaced on ex- plete carapace usually less than 6 mm, maximum but may be faintly developed at probably less than 9 mm. ternal surface, large, Cephalon subquadrate to transversely subrec- distal ends; median tubercle moderately tangular. Axial furrow well defined. Glabella with circular to oval base. Acrolobes uncon- bipartite (sag.) and slightly tapered forward; stricted to barely constricted. Border furrow length approximately two-thirds that of cephalon; moderately wide. Border with pair of postero- Je & Robison-Revision of a Trilobite Faunule 5

lateral spines, posterior section between spines based on the pygiclium described and figured as medially (sag.) widened and crescentiform. Baltagnostus (?) sp. by Lazarenko and Nikiforov Discussion.-On the basis of further study, (1968, p. 24, pl. 4, fig. 21), but that specimen and because of apparent gradational characters, appears to represent Ammagnostus or a related particularly among Manchurian specimens figured genus rather than Baltagnostus, and thus an un- by Endo and Resser (1937), Robison's (1964) equivocal occurrence of Baltagnostus in the Up- concept of Baltagnostus is here modified and ex- per Cambrian remains to be documented. panded to include species in which the pygidial In previous description and discussion, Robi- axis is separated from the posterior border furrow. son (1964, p. 525-527) indicated that the glabcllar Representatives of the genus in North America node of Baltagnostus had the form of a low, arc known to include B. centerensis (Resser, short, median ridge on the posterior end of the 1938), B. eurypyx Robison, 1964, B. marginalis glabella. Specimens of B. australis and B. dam esi (Rasetti, 1948), and at least two undescribed spe- exhibit such a posterior ridge as well as a typical, cies in Nevada and Utah. Proagnostus maryvil- though weak, median node near the midpoint of lensis Resser, 1938, is probably a synonym of B. the posterior glabellar lobe (Pl. 1, figs. 1, 2, 5). centerensis. A new species from Australia, B. Thus, the true glabellar node appears to be ef- australis, is added here. Agnostus rakuroensis faced on species such as B. eurypyx, and the Kobayashi, 1935, from South Korea was reas- posteromedial ridge appears to be a separate fea- signed to Peronopsis (Kobayashi, 1939, 1962), ture that characterizes at least B. australis, B. but is here transferred to Baltagnostus (see dis- damesi, and B. eurypyx. Because of either poor cussion of B. damesi). Species from Manchuria preservation or inadequate illustration of some include B. damesi (Resser & Endo in Endo & specimens, it is not possible at this time to deter- Resser, 1937), which may be a junior synonym mine whether or not a posteromedial glabellar of B. rakuroensis, and possibly some of the speci- ridge is a characteristic feature of all species of mens assigned to Agnostus comes by Resser and Baltagnostus. Endo (in Endo & Resser, 1937, pl. 30, fig. 17). As here redefined, Baltagnostus has an ob- Two species from Argentina, Baltagnostus served stratigraphie range through most of the hospitus Poulsen and B. mendozensis Poulsen, late Middle Cambrian (Ptychagnostus atavus to are more difficult to evaluate. The pygidium of Lejopyge calva assemblage-zones). Its occurrence B. mendozensis appears to lack the crescentiform in the early Middle Cambrian is questionable posterior border that is characteristic of Baltag- (see discussion of B. damesi). Representatives nostus, and its taxonomic assignment is further have been described from Australia (Queensland), hampered by lack of knowledge of the cephalon. Canada (Quebec), China (Manchuria, Shan- According to Poulsen (1960, p. 7-8), B. hospitus tung), North Korea, South Korea, and the United has confluent genae. the cephalon Further, ap- States (Alabama, Nevada, Tennessee, Texas, and pears to have a broadly rounded rather than a Utah). Reported occurrences in Argentina (Poul- constricted posterior glabella, and the pygidium sen, 1960) and Siberia (Lazarcnko & Nikiforov, lacks a prominent axial node as well as a cres- 1968) are probably in error. centic posterior border. Therefore, neither of these species may represent Baltagnostus. BALTAGNOSTUS AUSTRALIS Robison, n. sp. Opik (1967, p. 113) concluded that Baling- Plate 1, fi gures 3, 5-7, 9-11 nostus beltensis Lochman (in Lochman & Dun- can, 1944) probably belongs to Ammagnostus, Ettagnostns opinnts WHITEHOUSE (part), 1936, p. 87, pl. 8, fig. 12 (not figs. 10, 11). but could possibly represent Kormagnostus. I agree that the species should not be assigned to Holotype. -Cephalon, UQF69629 (PI. 1, fig. Baltagnostus, but have not further analyzed its 5). appropriate generic assignment. Other material.-Twenty-eight cephala and Baltagnostus sp. has been listed from the 27 pygidia. Upper Cambrian of the northwestern Siberian Description.-Cephalon subrectangular; length platform by Lazarenko and Datscnko (1967, p. and width of largest specimen 3.1 by 3.5 mm. 19, table 1). The entry in that list is probably Glabella distinctly narrower than adjacent genac 6 The University of Kansas Paleontological Contributions—Paper 90

(measured at position of transglabellar furrow); median ridge just above cephalic recess. Genae anterior lobe about one-third of glabellar length; of intermediate width. Lateral sides of acrolobes posterior lobe with weak median node at or slightly tapered toward anterior. slightly behind midpoint and a small median Thorax unknown. ridge just above cephalic recess. Genae wide. Pygidium subrectangular, length and width Lateral sides of acrolobes almost parallel or of lectotype 2.9 by 3.6 mm. Axis variable in slightly tapered toward anterior. length, reaching posterior border furrow in young Thorax unknown. holaspides, but usually separated from border Pygidium subrectangular; length and width furrow in older holaspides. Posterior axis taper- of largest specimen 2.7 by 3.1 mm. Axis rela- ing to slightly acute point. In specimens with tively short and well separated from posterior shorter axis, postaxial median furrow is wide border furrow; posterior part (behind second seg- and deep. Acrolobes slightly constricted. Border ment) tapering to slightly acute point. Postaxial furrow wide. median furrow usually deep and moderately wide. Discussion.—B. damesi is characterized by a Acrolobes unconstricted to barely constricted. glabella of intermediate width, a glabellar node Border furrow moderately wide and posterior part that is situated slightly anterior from the mid- may have anteromedial bend. point of the posterior lobe, indentation of the Discussion.—B. australis is characterized by posterior glabellar lobe by the basal lobes, and a its narrow glabella, relatively short pygidial axis, pygidial axis of intermediate and ontogenetically and well-developed postaxial median furrow. It variable length. It closely resembles and may be most closely resembles B. damesi (Resser & a junior synonym of B. rakuroensis, which Ko- Endo), but differs from that species by having a bayashi (1935, p. 103-104) originally assigned to narrow glabella, a median node situated at or Agnostus and later (1939, p. 189; 1962, p. 28-29) slightly behind the midpoint of the posterior reassigned to Perono psis. The partial preglabellar glabellar lobe, a shorter pygidial axis, and nar- median furrow, wide pygidial axis, crescentiform rower postaxial median furrow. posterior pygidial border and posterolateral border spines of rakuroensis are characteristic features of BALTAGNOSTUS DAMESI (Resser & Endo) Baltagnostus and the species is here assigned to Plate I, figures 1, 2, 4, 8 that genus. Kobayashi (1962, p. 29) noted the rignostus damesi RESSER & ENDO in ENDO & RESSER, 1937, similarity of damesi and rakuroensis, but distin- figs. KOBAYASHI, 1939, p. p. 157-158, pl. 30, 1-5; 579. guished them on the basis of the complete pre- Ettagnoctus damesi (Resser & Endo) WHITEHOUSE, 1939, glabellar median furrow that was erroneously p. 261; Lu and others, 1965, p. 51-52. Agnoctus haotungensis RESSER & ENDO in ENDO & RESSER, added to illustrations of damesi by Endo and 1937, p. 160, pl. 30, figs. 8-11. Resser (1937). All illustrated specimens of B. Ettagnostuç liaotungensis (Resser & Endo) WHITEHOUSE, rakuroensis are relatively small (young holas- 1939, p. 261. pides?) and rather poorly preserved. The cepha- Peronopsis liaotungensis (Resser & Endo) Ivslux, 1953, lon of at least one specimen of B. rakuroensis p. 11. (Kobayashi, 1962, pl. 3, fig. 1) has a glabella that AgnOftlIS ViatOr RESSLR & ENDO in ENDO & RESSER, 1937, p. 158-159, pl. 30, fig. 6. is considerably wider than is usual for B. damesi Ettagnostus via/or (Resser & Endo) WHITEnousE, 1939, and the pygidial axis appears to be relatively p. 261. broader and less tapered. Therefore, without ac- Peronopsis viator (Resser & Endo) IvsFrix, 1953, p. 12. cess to comparative material of B. rakuroensis, Lectotype.—Pygidium, USNM 57829a (Pl. 1, especially mature holaspicles, I tentatively retain fig. 4), designated here. B. damesi as a separate species. Description.—Cephalon subrectangular, length B. damesi also closely resembles B. australis and width of largest observed specimen 3.5 by n. sp., and the discussion of that species contains 3.9 mm. Glabella approximately equal in width a comparison of the two taxa. B. damesi can be to adjacent genae (measured at position of trans- most easily distinguished from North American glabellar furrow); anterior lobe about one-third species of Baltagnostus by its shorter pygidial glabellar length; posterior lobe with weak median axis, which is less tumid at the posterior end. node slightly anterior from midpoint, and tiny Three species, Agnostus damez-i, A. viator,

Jell & Robison—Revision of a Trilobite Faunule 7 and A. liaotungensis, were described by Endo and certainly known from rocks older than the middle Resser (1937) from the same locality of the Middle Cambrian (Ptychagnostus atavus Assem- Mapan Formation in Manchuria. Examination blage-zone). On the basis of evolutionary aspect, of the type specimens showed that cephalic illus- I suggest that B. damesi is probably of late rather trations of A. damesi (Endo & Resser, 1937, pl. than early Middle Cambrian age. Moreover, it 30, figs. 1, 2) have been erroneously retouched seems to be one of the undesignated species re- to show a complete preglabellar median furrow. ferred to by Endo for which former stratigraphie The same specimens are refigured here for com- placement "is no doubt in error." parison (Pl. 1, figs. 1, 2), and they show only a partial preglabellar median furrow as well as Genus DORYAGNOSTUS Kobayashi, 1939 other cephalic features that are characteristic of The generic concept of Doryagnostus is re- Baltagnostus. A. viator is based on a single small vised and discussed in an accompanying paper holaspid pygidium and A. liaotungensis is based (Robison, 1978). on a few relatively small specimens that together appear to represent no more than early holaspides DORYAGNOSTUS DELTOIDES Robison, n. sp. of damesi. Therefore, I consider the three spe- A. Plate 2, figures 7-9, 12 cies to be synonyms, and A. damesi, the senior syn- Enagnostus opimus WHITEHOUSE (part), 1936, p. 87, pl. onym, is transferred to Baltagnostus. The length 8, fig. 10 (not figs. 11, 12); HILL, PLAYFORD, & of the pygidial axis is somewhat variable, extend- WOODS, 1971, p. 18, pl. 9, fig. 4. ing almost to the posterior border furrow in young Holotype.—Cephalon UQF69624 (Pl. 2, fig. 9). holaspides, and becomes shorter with concomitant Other material.—One cephalon (UQF3194) appearance of a postaxial median furrow in older and two pygidia (UQF69625, 69626). holaspides. Description.—Cephalon subcircular; length A redescription of B. damesi is presented in by 4.6 mm, holotype this paper because of the erroneous concept of the and width of paratype 4.5 of intermediate width, species given by the retouched photographs ac- 4.7 by 5.0 mm. Glabella approximately equal to that of adjacent genae at companying the original description, and because level of transglabellar furrow; anterior part ogivi- the taxon is so similar to B. australis. White- point; position of house (1939, p. 261) also remarked on the simi- form with slight anteromedial on posterior lobe undetermined. larity of these forms and suggested that the con- median node Genae smooth. of intermediate cept of Euagnostus "should be restricted to such Border furrow types." What he did not recognize, however, is width, with well-developed deltoid area. that the holotype pygidium of the type species of Thorax unknown. Euagnostus is a representative of Perono psis, and Pygidium subcircular; length and width of application of the concept of Euagnostus to forms two known specimens 3.4 by 4.0 and 4.7 by 5.4 such as B. damesi is inappropriate. mm. Transaxial furrows effaced or only faintly Occurrence.—B. damesi has been reported developed near junction with axial furrow. Cen- from the Mapan Formation on Tschang-hsing-tao tral depression of posterior part weak or absent. (island southwest of Fu-chou, Pechili district), Posteromedial border furrow has slight forward Liao-tung, Manchuria. The Mapan Formation bow and posteromedial border slightly widened. was regarded by Endo (in Endo & Resser, 1937, Posterolateral border spines broken on both speci- p. 37-38) to be the basal Middle Cambrian mens; stubs of intermediate size. stratigraphie unit in southern Manchuria, and he Discussion.—D. deltoides is characterized by correlated its fauna with those of the lower Mid- its large, well-developed deltoid area (for defini- dle Cambrian "North America Ptarmigan and tion, see Robison, 1978) that extends almost to Langston formations in the Rocky Mountains." the glabella, and which is the basis for the species Endo (p. 38) further noted "that some mixture name. It is further characterized by a glabella of Mapan and Taitzu fossils exists in our collec- of intermediate width, a weak or absent median tions, particularly those made years ago, so that depression on the posterior part of the pygidial the placing of certain species is no doubt in error." axis, and a slight forward bow in the poster-0- Elsewhere in the world, Baltagnostus is not medial border furrow. 8 The University of Kansas Paleontological Contributions—Paper 90

Other recognized species of Doryagnostus arc Queensland. Its age is late Middle Cambrian D. incertus (Brtigger) and D. tvasatchensis Robi- (Ptychagnostus nathorsti Zone). son, which arc described and further discussed in an accompanying paper (Robison, 1978). D. HYPAGNOSTUS? sp. deltoides differs from both of those species by its Plate 2, figure 10 larger deltoid area. Its glabella is narrower and anteromedially more pointed than that of D. A single poorly preserved quadrate cephalon wasatchensis and broader than that of D. incertus. with effaced frontal glabellar lobe is present in In addition, its posterior pygidial axis is less cen- collection UQL256. Because of the tendency to- trally depressed and slightly less acuminate than ward effacement of the frontal glabellar lobe in that of D. incertus. Peronopsis op/mus, assignment to that species cannot be entirely ruled out. However, the quad- For a comparison with other agnostoids in rate cephalic outline is different from the more collection UQL256, refer to the discussion of subcircular outline of P. opimus. Also, in regard Perono psis opimus. to the posterior glabellar lobe, the lateral sides are more tapered, the anterolateral corners are Genus HYPAGNOSTUS Jaekel, 1909 more rounded, and the anterior furrow is more HYPAGNOSTUS cf. H. CLIPEUS Whitehouse deeply incised than in P. opimus. On the other Plate 2, figure 11 hand, the characters correspond closely to those of most species of Hypagnostus. Therefore, until Enagnostus sp. WurrntousE, 1939, 260-261. P. more material becomes available for comparison, Discussion.—In his documentation of trilo- the specimen is questionably assigned to Hy- bites from northwestern Australia, Whitehouse pagnostus. (1939, p. 260-261) referred one pygidium to The specimen differs from H. parvifrons, of Euagnostus sp., but did not figure it. That speci- similar age, by its quadrate outline, and its glabella men is here illustrated for the first time. It is is more than half the cephalic length. characterized by an unfurrowed axis that is nearly parallel sided in the anterior two-thirds of its Genus PERONOPSIS Corda length and tapers in the posterior one-third to a Peronopsis CORDA in HAWLE & CORDA, 1847, p. 115; slightly acute point. The axial tubercle is weak; ROBISON, 1964, p. 529-530 (for additional and nearly the postaxial furrow is short and wide, and complete synonymy to 1964); 45unc, 1967, p. 75; merges with a posteromedially widened border PALMER, 1968, p. 31; PER & VANEK, 1971, p. 269- furrow; and the border widens appreciably from 270; PALMER & GATEHOUSE, 1972, p. 9; JAco, 1976, p. 136. anterior to posterior, and is nonspinose. All of Ettagnostus WHITEHOUSE, 1936, p. 87; HOWELL, 1959, p. these characters are typical of some species of 184; Lu and others, 1965, p. 51; (lipm, 1967, p. 78. Hypagnostus. NEW SYNONYMY. The pygidium and a large complete specimen Type species.—Battus integer BEYRICH, 1845, of Doryagnostus incertus (Robison, 1978, pl. 2, p. 44, by monotypy. fig. 8) are preserved together on a piece of lime- Diagnosis.—Peronopsis is characterized by a stone, and are from the same locality as the bipartite glabella with simple basal lobes and ab- holotype of Hypagnostus clipeus Whitehouse. By sence of a preglabellar median furrow. The axial comparison, figured pygidia of H. clipeus (White- furrow is generally well defined, but rarely may house, 1939, pl. 25, figs. 25, 26; Hill, Playford & be shallow around the anterior glabella; genae Woods, 1971, pl. 9, fig. 5) are somewhat crushed and pleural fields are usually smooth; and trans- and appear to have a narrower posterior border. verse furrows of the pygidial axis are commonly Therefore, taxonomic assignment of this pygidium effaced or only poorly developed. Characters such is questionable, but it appears likely to be a as shape and segmentation of the pygidial axis, variant individual of H. clipeus. position of the glabellar node, width of border Occurrence.—One pygidium, UQF69639, from furrows and borders, and presence or absence of the V Creek Limestone at the road crossing of pygidial border spines or swellings vary within the V Creek between "Undilla" and Thorntonia, genus, but tend to be fairly stable within given

Jell & Robison—Revision of a Trilobite Faunule 9 populations. Therefore, those kinds of features described as Doryagnostus tvasatchensis in an are considered to be of value for differentiation accompanying paper (Robison, 1978). of species. Other characters, such as outlines of All of the agnostoids from Whitehouse's col- the cephalon and pygidium, relative size of the lection UQL256 are redescribcd in this paper, and axial lobe, and presence or absence of a postaxial the holotype of E. opimus is now assigned to median furrow, may vary during ontogeny. Thus, Perono psis, whereas the two figured paratypcs usually less taxonomic importance is attached to are reassigned to Baltagnostus australis, n. sp., those kinds of features. and Doryagnostus deltoides, n. sp. Because of Discussion.—A detailed analysis and discus- the change in generic assignment of the holotype sion of Peronopsis is in preparation for separate of its type species, Euagnostus is suppressed as a publication by Robison. For purposes of this subjective junior synonym of Peronopsis. Addi- paper, only the new synonomy of Euagnostus is tional documentation and support for these ac- considered. tions are contained in the separate discussions of Whitehouse (1936, p. 87) described Euagnos- each of the species mentioned. tus as a new genus and designated E. opimus as its type species. He also figured and assigned PERONOPSIS OPIMUS (Whitehouse) three specimens (one ccphalon and two pygidia) Plate 1, figures 1-6 to that species. In a later brief discussion, Ôpik Euagnostus opimus WHITEHOUSE (part), 1936, p. 87, pl. (1967, p. 78) noted that "the holotype of E. 8, fig. 11 (not figs. 10, 12). opimus is an imperfect pygidium which recalls a Hypagnostus; the associated cephalon, however, Holotype.—Pygidium, UQF3195 (Whitehouse, may belong to a species related to Ptychagnostus 1936, pl. 8, fig. 11; refigured here Pl. 1, fig. 4). con vexus Westergaard or P. stenorhachis Other material.—Ten cephala and three py- wall." In concluding the same discussion, he also gidia in collection UQL256. noted that "it is still not impossible that the holo- Description.—Cephalon subcircular, width type pygidium and the cephalon arc conspecific." about 10 percent greater than length; maximum Subsequently, and without further taxonomic re- observed length 4.8 mm. Axial furrow moder- mark, Opik (1970, p. 4) proposed substitution ately developed posteriorly, weak or nearly ef- of Euagnostus opimus for Hypagnostus parvifrons faced around anterior glabellar lobe. Glabella as a zonal name in Australia because H. parvi- slightly tapered; anterior lobe low, and may have Irons was unknown in that country. The name, obtuse anteromcdian point; posterior lobe tumid, Euagnostus opimus Zone, has been cited in a with faint axial node situated well forward from number of succeeding Australian publications. midpoint. Border furrow narrow. Border narrow In 1971, Hill, Playford, and Woods (p. 18, pl. 9, to moderately wide. fig. 4) refigured the paratype cephalon of E. Thorax unknown. opimus and gave an abbreviated description of Pygidium subcircular to subrectangular, width that specimen. about 15 to 20 percent greater than length; maxi- During 1972, Robison discovered Euagnostus- mum observed length (holotypc) 3.9 mm. Axis like trilobites in the middle Middle Cambrian of tapering unevenly to acute point, extending al- northern Utah. On the basis of available descrip- most to posterior border furrow; transaxial fur- tions and illustrations, those specimens were ten- rows effaced; median tubercle slightly elongate tatively identified as Euagnostus opimus and have (sag.), of moderate size. Postaxial median furrow been mentioned as such in one faunal list (Robi- short and wide; almost eliminated by near merger son, 1976, p. 105). An opportunity to examine of axial and border furrows. Border furrow nar- Whitehouse's collection UOL256 during the sum- row to moderately wide. Border narrow anteri- mer of 1976 led to the conclusion that the three orly, widening appreciably toward posterior; with specimens illustrated and assigned to E. opimus obvious posterolateral swellings, but with true by Whitehouse (1936) actually represent three spines. species each belonging to a different genus. More- Discussion.—P. opiums is characterized by the over, it is further concluded that the specimens weak or nearly effaced axial furrow around the from Utah represent a new species, which is anterior glabellar lobe, anterior position of its

10 The University of Kansas Paleontological Contributions—Paper 90 glabellar node, tapered and elongate pygidial axis The pygidial axis of B. australis is more tumid with effaced transaxial furrows, and a pair of immediately behind the second segment than are posterolateral swellings on the pygidial border. the other two species, and D. deltoides has an ap- Of the many species of Peronopsis, P. opimus preciably longer postaxial median furrow than most closely resembles P. interstricta (cf. Robison, does P. op/mus or B. austral/s. 1964, pl. 82, figs. 1-15, 18). These species can be Near effacement of the anterior axial furrow distinguished fairly easily, however, by the more of P. opimus results in a form that is transitional effaced anterior axial furrow, more anteriorly between Peronopsis and Hypagnostus. Neverthe- situated glabellar node, and more elongate pygidial less, because its stratigraphie occurrence is con- axis and shorter postaxial median furrow of P. siderably higher than the lowest known occur- opimus. Also, the glabella of P. opimus tends rence of species of Hypagnostus, P. op/mus can to have a slight anteromedian point, whereas such be excluded as a possible ancestor of Hypagnostus. a feature is usually not present on P. interstricta. The form does, however, provide an indication P. opimus also closely resembles some speci- of possible homeomorphy among species that have mens of P. scutalis that were illustrated by Wester- been assigned to Hypagnostus. Ord (1946, pl. 4, figs. 4-11), but differs from that species by its more effaced anterior axial Eodiscoid Trilobite furrow, more anteriorly situated glabellar node, [by P. A. Jell] broader pygidium, and prominent pygidial border swellings, which are only faint or absent on P. Family EODISCIDAE Raymond, 1913 scutalis. On the basis of a bimodal difference in Genus OPSIDISCUS Westergard, 1949 axial length on the pygidiurn, it appears likely that two species may actually be included within OPSIDISCUS MICROSPINUS Jell the P. scutalis of Westergard. Plate 4, fi gures 8a, b P. bfighamensis is one of the few other species Opsidiccus microspinus JELL, 1975, p. 80, pl. 24, figs. 1, 2, 4; pl. 26, figs. 3, 4, 7, 8. of Peronopsis characterized by a pair of swellings on the posterolateral pygidial border; however, P. Material.—One cephalon in collection UQL256. brighamensis differs from P. op/mus by its nar- Remarks.—The almost unsegmented and an- rower glabella, less conical pygidial axis, moder- teriorly pointed glabella, weak border scrobicules, ately- to well-defined transaxial furrows, and well-impressed border furrow, and steep promi- larger median tubercle on the pygidium. nent facets are indicative of O. microspinus. Whitehouse (1936) failed to recognize differ- Spaces for the eyes are clearly exhibited high on ences between forms in collection UQL256 that the cheeks, and facial sutures are absent. Dam- are here assigned to Peronopsis opimus (cf. Pl. 2, age to the occipital spine and lack of a pygidium fig. 4a-c), Doryagnostus dc/to/des (cf. Pl. 2, fig. prevent conclusive taxonomic assignment; how- 7a, b), and Baltagnostus australis (cf. PI. 1, fig. ever, available features leave little doubt about 9a, b); however, these species differ in many the identity of this cephalon. characters. Cephala can be most readily distin- guished by the lack of a preglabellar median fur- Polymeroid Trilobites row in P. opimus, the presence of an incomplete [by P. A. jell] preglabellar median furrow that is best developed posteriorly in B. australis, and the presence of a Family ANOMOCARIDAE Poulsen, 1927 complete preglabellar median furrow and dis- Opik (1967, p. 212) removed the genera tinctive deltoid area in D. deltoides. Also, the Westergaardella Kobayashi, 1962, Chondranomo- glabella is relatively broader in P. opin2us and care Polctaeva in Chernysheva and others, 1956, D. dc/to/des than in B. australis, and the anterior Pseudanomocarina Chernysheva in Chernysheva axial furrow is less well defined in P. op/inns and others, 1956, and Schoriella Sivov in Khalfin, than in D. deltoides and B. austral/s. On the 1955, from the Anomocaridae. These, along with pygidial border, D. dc/to/des and B. australis each Auritama Opik, 1967, were placed in a new fam- have a pair of posterolateral spines, whereas P. ily, Auritamidae, which Opik diagnosed as "Ano- op/mus has only a corresponding pair of swellings. mocaracea with a long parallel-sided glabella and Jell & Robison—Revision of a Trilobite Faunale 11

a relatively small pygidium." None of these gen- slightly forward, rounded-angular or rounded an- era are known from articulated specimens, and teriorly, weakly inflated, having a weak median therefore relative pygidial size is a poor diagnostic longitudinal keel. Glabellar furrows usually ab- character. Of the two Middle Cambrian genera, sent. In some species one or two pairs of very Pseudanomocarina is here considered to be a indistinct, broad, shallow furrows are outlined synonym of Chondranomocare, and all species of on the glabella. Pseudanom ocarina except the type, P. plana, arc Fixed cheeks narrow. Palpebral lobe [eyelid] assigned to a new genus Sudanomocarina, which quite large, with an angular break toward the along with Chondnmomocare, contains several rear of its length. Eye surface not present. species in which the glabella is not long (i.e., Frontal limb quite wide, with a moderately prom- does not reach the border furrow) and it tapers inent, low terrace extending parallel to the an- anteriorly. Pygidial characters of these two genera terior margin of the cranichum and situated at are also quite distinct from those of Auritanta, different distances from the glabella in different particularly in the lack of spines. The genus species. Anterior border furrow distinct or vague. Schoriella is based on the early Late Cambrian Border narrow, flat, slightly upturned. Facial species S. schorica Sivov (in Khalfin, 1955) from suture in front of eye diverging in bow-shaped southern Siberia, which superficially resembles curve as it extends to the anterior margin. some species of Chondranomocare, but the possi- Pygidium slightly drawn out in width, having bility of homeomorphy makes the relationship of weakly concave posterior margin. Axis high, these species uncertain. Most other species pre- rounded posteriorly, having longitudinal keel on viously referred to Schoriclla are more correctly the posterior margin. On the axis and on the placed in the genus Schoriecare Rozova, 1964. flat pleural areas, well-impressed furrows produce Close relationship may exist between the other undulating rounded curves on the segmental cross two Late Cambrian genera, Auritama and Wester- section. gaardella, but Schoriella is certainly distinct from Discussion.—Chondranomocare was not com- these with respect to its anteriorly tapered glabella pared with other genera at the time of its erec- and palpebral lobes close to the glabella. There- tion by Poletaeva (in Chernysheva and others, fore, I cannot accept the grouping of these five 1956), and only Chernysheva (1961, p. 197) and genera into a separate family. Moreover, if the Palmer and Gatehouse (1972, p. 21) have dis- Auritamidae is to be accepted, a better diagnosis cussed it subsequently. Chernysheva recognized must be provided. I retain Chondanomocare similarities between Chondranomocare and Psett- (= Pseudanomocarina), the new genus Sudan- danomocarina in glabellar outline and segmenta- 0m ocarina, and tentatively Schoriella, in the tion, in dimensions and form of the fixigenae, Anomocaridae even though its taxonomy is poorly and in dimensions and position of the palpebral understood and a worldwide review is long over- lobes. The only distinctive difference that she due. noted is the structure of the preglabellar part of the craniclium. Genus CHONDRANOMOCARE Poletaeva One important feature—glabellar shape—was Chondranomocare POLETAEVA in CHERNYSHEVA and others, mentioned in the original diagnosis of Pseudano- 1956, p. 169; EGOROVA in KHALFIN, 1960, p. 214; mocarina as being rectangular or just slightly CHERNYSHEVA, 1961, p. 196; PALMER & GATEHOUSE, 1972, p. 21. tapered forward, and this has been incorporated Psettdanomotarina CHERNYSHEVA in CHERNYSHEVA and into the generic concept (not based on the type others, 1956, p. 166; CHERNYSHEVA, 1961, p. 187. species) by subsequent Russian authors. In com- NEW SYNONYMY. bination, the rectangular glabella and very short Type species.—Chondranomocare bidiensis or absent preglabellar field distinguish a group POLETAEVA in Chernysheva and others, 1956, p. of Russian species and one new Australian species 170, by original designation. that I refer to the new genus Sudanomocarina. Diagnosis (translated from Poletaeva in Cher- The holotype of the type species of Pseudanomo- nysheva and others, 1956, p. 169, by P.A.J.).— carina, P. plana, lacks this combination of char- Trilobites of less than average size. Cephalon acters. Furthermore, because of its rounded not much larger than pygidium. Glabella tapers glabellar anterior, tropidium behind a very shal- 12 The University of Kansas Paleontological Contributions-Paper 90 low border furrow, and similar frontal area, P. its palpebral lobes terminate some distance from plana is here considered to belong to Chondrano- the axial furrow, both anteriorly and posteriorly, mocare and to be fairly closely related to C. and are more evenly rounded; the frontal area bidjensis. Comparison of the holotype of P. lacks a terrace; the long genal spine is equal in plana (Chcrnysheva, 1961, pl. 22, figs. la, b) length to the remainder of the librigena; the with the remaining cranidia assigned to the spe- pygidial margin is well rounded; and the trans- cies (Chernysheva, 1961, pl. 22, figs. 2-8) clearly verse pleural and interpleural furrows do not ab- illustrates these significant differences. axially swing strongly toward the posterior. Su- Palmer and Gatehouse (1972, p. 21) stated danomocarina is more difficult to distinguish that "the concave rather than flat or convex from Chondranomocare, and the pygidia are in- border distinguishes Chondranomocare from Pseu- distinguishable. Cranidia of Sudanomocarina danonlocarina," but the original diagnoses stated have no terrace on the preglabellar field. Indeed, that Chondranomocare has a flat border whereas there is virtually no preglabellar field, as the Psezidanomocarina has a convex or concave one. glabella reaches almost to the border furrow and I do not consider the shape of the border's lateral is more quadrate and proportionally wider. profile to be a diagnostic feature of either genus. Two other genera that deserve mention are If visible, lateral glabellar furrows consist of Schoriella and Schoriecare. As already mentioned, three pairs in species of Chondranomocare: 1P the Late Cambrian Schoriella schorica is a pos- divide adaxially and are directed strongly pos- sible homeomorph of the Middle Cambrian Chon- teriorly; 2P and 3P are narrower and more trans- dranomocare con fertum, and it can be distin- verse. The terrace (or tropidium) on the pre- guished only by its better impressed border glabellar field (frontal limb of Poletaeva's diag- furrow and more strongly tapered anterior gla- nosis) is a variable feature. It is present on most bella, which also appears to taper posteriorly. S. figured specimens, but some do not appear to ex- schorica is known from a single figured cra- hibit the feature (e.g., Chernysheva, 1961, pl. 25, nidium, however, and is too poorly known to fig. 8; Egorova in Egorova & Savitsky, 1969, pl. warrant more comment. Like C. con fertum, and 38, figs. 17, 18; pl. 40, fig. 2; Palmer & Gatehouse, apparently Schoriella, Schoriecare has an extreme 1972, pl. 3, figs. 18, 19). posterior extension of the palpebral lobe, being On the originally figured pygidium of Chon- distinguished by its much longer and more ex- dranomocare bid jensis (Chernysheva and others, panded frontal area and its pygidial characters. 1956, pl. 31, fig. 4) the axis is extremely short, Occurrence.-Chondranomocare is known but on other figured pygidia (C. eminens Cherny- from the late Amga Stage of Siberia and from sheva, 1961, pl. 25, fig. 10; C. exilis Egorova in equivalent strata of the Hypagnostus parvifrons Egorova & Savitsky, 1969, pl. 40, fig. 7; C. spe- Zone in northwestern Queensland. Age of C. ciosum M. Romanenko (figured by Egorova, australis in Antarctica is uncertain, but is said to 1969, pl. 39, fig. 20); C. bucculentum Lazarenko, be early late Middle Cambrian, and may be con- 1965, pl. 1, fig. 18; C. australis Palmer & Gate- temporaneous with C. con fertum from Australia. house, 1972, pl. 3, fig. 24) it is proportionally longer. Another consistent feature of the py- CHONDRANOMOCARE CONFERTUM gidium is the pair of angularities in the margin (Whitehouse) adjacent to the abaxial ends of the anterior pleural Figure 2; Plate 3, figures 1-19 furrows. Distinguishing Chondranomocare from other Anornocare confertum WHITEHOUSE, 1939, p. 223, pl. 23, figs. 22-28. genera in the family is easy at present, but I ex- pect that a full review of the family will show Holotype.-Cranidium UQF3354 (White- considerable intrageneric variation, making ge- house, 1939, pl. 23, fig. 22; refigured here, pl. 3, neric definition more difficult. Two genera are fig. 1 1 a-c). mentioned in this regard. Anon2ocare Angelin, Other material.-In the type collection are 1854, based on A. laeve (Angelin, 1851) from an additional 12 cranidia, 4 librigenae, and 8 the late Middle Cambrian Andrarum Limestone pygidia, including the figured specimens. I have of Sweden, has several distinguishing features: collected the species from contemporaneous strata Jell & Robison—Revision of a Trilobite Faunule 13 at several other localities in the Undilla basin near midlength; posterior margin almost straight ex- Thorntonia homestead, and it is present in several cept for slight anterior curve near axial furrow. undescribed collections from the same area made Axial furrow for most of its length barely more by F. W. Whitehouse and deposited at the Uni- than a change in slope from glabella to genae, versity of Queensland. poorly impressed posteriorly. Low para frontal Diagnosis.—Cranidium as long as wide, trans- band across front of glahella with extremely faint verse profile weakly convex, but lateral profile bounding furrows. strongly downturned in anterior part; ornament Anterior fixigenae strongly expanded into tri- finely granulose. Glabella 0.8 cranidial length, angular areas abaxially (Pl. 3, fig. 11), sometimes tapering slightly forward, sides straight, antero- of almost uniform length (PI. 3, fig. 13). Border lateral corners somewhat angular, anterior margin furrow very shallow; in some specimens postero- nearly straight to slightly convex; three pairs of medially expanded with slight pits immediately lateral furrows poorly developed, midline with a lateral from expansion, pits possibly relating to weak but distinct longitudinal keel. Preglabellar slight anterior convexities in terrace on preglabel- field short, 0.1 cephalic length, with single trans- lar field. Border of near constant width except verse terrace (not visible on all specimens) near laterally where cut by oblique facial suture. border furrow. Anterior border flat to slightly Palpebral furrow poorly impressed throughout, convex, upturned. Palpebral area of fixigena nar- weakest near midlength, roughly paralleling outer row, flat to slightly convex, widest adjacent to margin of palpebral lobe, curving posteriorly into furrow IP and tapered to anterior point; palpe- border furrow. Palpebral lobe very slightly con- bral lobe wide (half width of fixigena at mid- vex (tr.), slightly wider posteriorly; in a few point), long (0.6 cephalic length), joined to specimens with faint indication of a furrow bi- anterior glabellar lobe, straight anteriorly then secting it lengthwise. strongly curved to termination opposite mid- Librigena with very low but distinct eye socle. length of occipital ring; posterior area of fixigenae Subocular area with faint genal caeca, narrowest wide, short, well beneath posterior of palpebral anteriorly, expanding to posterior border furrow. lobe, angled slightly down. Anterior facial sutures Border 0.35 of total width anteriorly, weakly con- diverging strongly from points near glabella. vex (tr.), greatly expanded at base of genal spine, Librigena relatively flat with shallow border fur- then tapering strongly toward axis on short pos- row, strong genal spine, and steep to vertical terior part. Facial suture runs obliquely across posteroproximal extremity. the border furrow and posterior border to pos- Pygidium moderately convex. Axis 0.75 py- terior margin. Extremely faint epiborder furrow gidial length, with four poorly defined ring fur- apparent on posterior edge just adaxial from rows. Pleural furrows transverse adaxially but genal spine. Genal spine flat to slightly convex, exsagittal abaxially (i.e., with sharp bend near with broad base, tapering strongly on proximal midlength). Margin weakly angulatc adjacent to third, continuing even curve of cephalic margin. termination of anterior pleural furrows, slightly Border furrow slightly wider, deeper, more U- indented behind axis. Pleural furrows extend shaped in section toward posterior. almost to margin. Shallow furrow just adaxial Hypostoma convex with stout anterior wings; from inner doublural margin. border anteriorly short, strongly upturned, taper- Description.—Lateral glabellar furrows very ing strongly toward posterior; lateral border fur- poorly impressed, defined by lack of ornamenta- row broad and shallow anteriorly, narrowing tion; 1P directed back at 45° from sagittal line, posteriorly. Ornamentation on main body con- long, with marked curve just inside axial furrow; sisting of large low tubercles. 3P transverse, with marked elongate transverse Thoracic segments short with strongly curved antcromedian muscle scar and slight anterior and pointed pleural tips. Axial ring convex with constriction adjacent to palpebral lobe. Occipital short (0.2 length) articulating half ring, moder- furrow mostly well impressed with gently sloping ately impressed articulating furrow, and long walls, but extremely steep near axial furrow; flat articulating lobe that is slightly longer ab- lateral muscle scars prominent. Occipital ring axially. Pleural furrow moderately impressed, flat (sag.), of constant length, with tubercle at with slight pit just beyond fulcrum, anterior wall

14 The University of Kansas Paleontological Contributions—Paper 90 steep and posterior wall gentle; extending from relative length of the glabella and its separation anterior of segment at axial furrow to midlength from the anterior margin, a migration of the at fulcrum, terminating well down pleural spine facial suture closer to the glabella, a narrowing near posterior margin. of the anterior cranidium, a transition from a Pygidium with poorly defined axial furrow, rectangular to anteriorly tapering glabella, and steep slope from rear of axis to border, short a narrowing of the fixigenae. articulating half ring, and articulating furrow no better developed than ring furrows, articulating Genus SUDANOMOCARINA Jell, n. gen. facets not apparent on available material. Axis Type species.—Sudanomocarina changi JELL, tapering slightly toward posterior, terminus n. sp. rounded with two large rounded knobs. Low Other included species.—Pseudanomocarina sagittal ridge extending from posterior axis al- acutata Bognibova in Bognibova and others, most to margin. Faint furrow just anterior from 1971; P. aojiformis Chernysheva in Chernysheva first ring furrow separates a structure equal in and others, 1956; P. be/la Hajrullina in Repina area and shape to articulating half ring. Pleural and others, 1975; P. eldachica Bognibova in furrows moderately deep, extending almost to Bognibova and others, 1971; P. horrida Cherny- margin at anterior, very weak and not extending sheva, 1961; P. parva Chernysheva, 1961; P. laterally beyond inner margin of doublure at tabatica Repina in Khalfin, 1960. Except for the posterior. Doublure narrow anteriorly and pos- holotype, all specimens referred to P. plana by teriorly, extremely wide (0.8 axial width) in be- Chernysheva (1961, pl. 22, figs. 2-10) belong to tween; very closely oppressed beneath dorsal exo- Sudanonlocarina, but need a new species name. skeleton. A faint but distinct furrow just inside Diagnosis.—Cranidium weakly convex; gla- inner margin of the doublure terminates at right bella rectangular or slightly tapered, lateral fur- angles to anterior pleural furrow, and a broad rows absent or very weakly impressed, occipital shallow border furrow is situated abaxially to it. ring flat and lengthened medially, with or with- Remarks.—Chondranomocare con fertum is a out pointed median tubercle; palpebral areas of distinctive species separated from all others in fixigenae very narrow (less than 0.33 glabellar the genus by the termination of its palpebral lobes width), weakly convex; palpebral lobe wide, opposite the occipital ring. It is perhaps most arcuate, situated on posterior half of cranidium, similar to C. exilis Egorova (in Egorova & about half length of glabella; preglabellar field Savitsky, 1969), but its anterior border furrow short or absent; anterior border weakly convex is less distinct, its anterior border is less upturned, or concave; anterior facial sutures diverging in it is narrower between facial sutures at the an- bow-shaped curve. Librigena convex with long terior palpebral lobes, and its pygidium is more spine. quadrate. Thorax unknown. Although C. con fertum could be removed to Pygidium transverse with curved anterior a monotypic genus, I refrain from such action border, sometimes with a slight indentation on because the species is probably part of a lineage posterior margin; greatest width near midlength; arising from Chondranomocare, and until this axis not reaching posterior border, generally with lineage is better understood, it is pointless to 5 or 6 poorly defined axial rings; pleural areas create a new generic name. With regard to weakly convex toward axis, but weakly concave variation within the genus, it should be kept in toward border; pleural ribs distinctly turned to- mind that C. con fertum is on the periphery. ward posterior near midwidth, distinctly divided The specimen described by Egorova (in by interpleural furrow; surface smooth. Egorova & Savitsky, 1969, p. 209, pl. 40, fig. 1) Discussion.—Sudanomocarina is erected to re- as Stella sp. should probably be referred to C. place the name Pseudanomocarina because the exilis. It also has librigenae almost identical with holotype of the type species of Pseudanon2ocarina, those of C. con fertum. P. plana, is transferred to Chondranomocare. Morphogeny.—By means of coordinate trans- Other species that have previously been assigned formation (Fig. 2), four growth stages of C. to Pseudanomocarina are here transferred to conferturn are used to illustrate a reduction in Sudanom ocarina.

Jell & Robison—Revision of a Trilobite Faunule 15

A close relationship between Sudanomocarina changi, in the Peronopsis opimus Zone of north- and Chondranomocare is apparent and their geo- western Queensland. graphic and stratigraphic association is compatible with this conclusion. Presence or absence of a SUDANOMOCARINA CHANGI Jell, n. sp. tropidium has apparently been used by some Rus- Plate 4, figures 9-16 sian workers to distinguish species of Chondrano- mocare from species formerly assigned to Pseu- Holotype.—Cranidium UQF69375 from UQL danomocarina; however, variability of that feature 463 (Pl. 4, fig. 14). in Chondranomocare confertum weakens its diag- Other material.—Two badly damaged cranidia nostic value. and 1 damaged pygidium from UQL256, and 5 Occurrence.—Sudanomocarina ranges through- cranidia from UQL463. out the Amga Stage on the Siberian platform Diagnosis.—Sudanomocarina characterized by and in the Sayan Altay region where it also very slightly tapered glabella with rounded an- ranges up into the lower Maya Stage (Bognibova terolateral corners, blunt anterior, and extremely & Shcheglov, 1971). Its only other occurrence weak lateral glabellar furrows; frontal area with recorded to date is that of the type species, S. uniformly short, flat to slightly convex border,

A

A n1111111Milirr 1.10t*- WOMIIRWIN odIALIONION% D

D FIG, 2. Proportional changes in cranidial morphogcny of Chondranomocare confertum interpreted by coordinate transformation; A drawn from specimen on Plate 3, figure 2 ( X73); B from Plate 3, figure 3 (X46); C from Plate 3, figure 5 (X11); and D from Plate 3, figure 13 (X4).

16 The University of Kansas Paleontological Contributions—Paper 90 very short to absent preglabellar field; fixigenac margin. Lateral margin well rounded into lobes narrow; palpebral lobes long and weakly convex with marked posteromedian indentation. Axis with anterior ends close to glabella; anterior 0.7 sagittal length, tapered slightly toward pos- facial sutures moderately divergent. Pygidium terior; half ring very short; two straight, mod- about twice as wide as long, with rounded erately deep ring furrows define relatively short weakly lobate pleural margins, posteromedian rings; terminal piece large, posteriorly rounded, marginal indentation, wide doublure. 0.6 sagittal length. Anterior axial ring marked Description.—Cranidium almost as wide as by posteromedian excavation. Axial furrow in- long, only slightly convex, but anterior glabella dicated by change in slope from axis to pleural slopes moderately forward to border furrow. area. Pleural regions sloping gently to margin, Glabella with straight, almost parallel sides, with three pleural furrows decreasing in depth straight anterior margin across sagittal line, posteriorly and one faint interpleural furrow. curved anterolateral corners, and three pairs of Anterior border furrow (or first pleural furrow) weak lateral glabellar furrows in some specimens well impressed parallel to anterior and lateral (Pl. 4, figs. 10b, 12). Furrow 1P directed toward margins, exsagittal in distal portion. Doublure posterior and weakly bifurcate adaxially, 2P trans- wide posterolaterally (0.5 width of pleural area), verse, 3P almost imperceptible and directed an- tapering anteriorly and adaxially. teriorly. Occipital furrow with steep anterior wall Dorsal surface of exoskeleton finely papillose. and slight apodemal pits just abaxial from mid- Remarks.—This new species closely resembles point between sagittal line and axial furrow. Oc- most specimens figured by Chernysheva (1961, cipital ring short, tapering laterally to half sagittal pl. 22, figs. 3-8) as Pseudanomocarina plana, espe- length, with small median node at midlength. cially her figure 8. In fact, it is impossible to Preglabellar field extremely short or absent. An- distinguish most cranidia from the Soviet Union terior cranidium moderately downsloping, slope and Queensland. The holotype of P. plana, how- increasing anterolaterally. Border furrow poorly ever, can be differentiated by its larger frontal impressed but distinct, corresponding to marked area, longer anterior border, more rounded glabel- change of slope, width constant (sag.), depth lar anterior, and slightly shorter eyes, and is here almost constant. Border weakly convex, narrow- transferred to Chondranomocare bid jensis. Two ing slightly toward lateral ends, horizontal to pygidia figured by Chernysheva (1961, pl. 22, gently arched in transverse profile. Palpebral figs. 9, 10) are also distinct from the Queensland area of fixigena very narrow (0.30 to 0.45 basal specimens. Other material figured by her in 1961 glabellar width), weakly convex, variable in shape, is probably conspecific with S. changi from widest point at or behind midlength. Palpebral Queensland, but is not formally designated as lobe 0.40 to 0.45 glabellar length, sloping down such in the absence of adequate figures or access abaxially, narrowing slightly toward posterior. to Soviet material. S. aojiformis has a parallel- Eye ridge short but usually prominent. Palpebral sided and proportionally larger glabella, more furrow shallow but distinct, swinging abaxially gently sloping glabellar anterior, and more tri- behind eye, shallowest near midlength. Facial angular pygidium. Other species (e.g., S. parva sutures diverging anteriorly at 15° from exsagittal and S. eldachica) are distinguished by a longer lines, adaxial points (anterior and posterior to preglabellar field, straight- to parallel-sided gla- palpebral lobe) close to axial furrow and in the bella, and deeper lateral glabellar furrows. but pos- same exsagittal line in most specimens, The two specimens from UQL256 are imma- more terior adaxial point may be slightly abaxial. ture and the species description is based largely to Posterior area of fixigenae 0.15 0.20 (exsag.) on the material from UQL463. glabellar length, extending further abaxially than palpebral lobe, steeply downsloping abaxially. Family DAMESELLIDAE Kobayashi, 1935 Posterior border furrow well impressed behind midlength, with gentle forward arch. Genus PRODAMESELLA Chang

Pygidium reniform, width 1.5 times length, Produniesclla CHANG, 1959, p. 196, 218; KOBAYASHI, 1960, with low posterior convexity, moderate lateral p. 352; Ix and others, 1965, p. 397 [nom. nod., convexity, and a steep slope from axis to posterior CHANG, 1957, pl. 1, fig. 5]. Jell & Robison—Revision of a Trilobite Faunule 17

Type species.—Prodamesella con vexa CHANG, anteriorly and adjacent to occipital ring. Glabella 1959, p. 196, by original designation. about one-third cranidial width, reaching anterior Diagnosis.—Chang's diagnosis (1959, p. 196; border furrow, tapering slightly forward to 3P, p. 218 for English translation) is followed here. slightly expanded in frontal lobe that has slight Discussion.—The two species, Damesella anteromcdian indentation and bibbed appear- quadrata Resser and Endo (in Endo & Resser, ance. Three pairs of lateral glabellar furrows 1937) and Olenoides- manchuriensis Endo, 1944, laterally deep and quite narrow; 1P and 2P ex- tentatively referred to Prodamesella by Chang tending adaxially and slightly to posterior from (1959, p. 219) are here excluded from that genus, axial furrow, 3P almost transverse. Occipital although their correct generic placement is un- furrow well impressed with slight lateral pits; certain. D. quadrata has a more quadrate glabella, directed slightly posterior from axial furrow to much longer palpebral lobes, and wider posterior pits, then anterior in gentle arch. Occipital ring fixigenae than P. convexa. Short eyes are char- 0.2 glabellar length, with strongly convex pos- acteristic of the Damesellidae and D. quadrata terior margin, strong lateral taper; four tiny pits can be excluded on that feature alone. O. man- surrounding a small posteromedian tubercle on churiensis is distinguished by its more quadrate a faintly raised circular area extending full length glabella with poorly impressed lateral glabellar of ring, anterior pair of pits more widely sepa- furrows, convex anterior border, wider fixigenae, rated (tr.) than posterior pair. Anterior border and more quadrate cranidium. Nature of the furrow U-shaped in section, slightly expanded in anterior border and glabella are of generic sig- front of glabella where posterior wall is convex. nificance. Anterior border flat but strongly upturned, slightly Prodamesella is here restricted to P. con vexa expanded posteromedially, tapered laterally. Fixi- and P. biserrata, and is defined by the truncato- genae convex and narrower toward anterior, conical glabella with deeply incised lateral glabel- crossed by high prominent eye ridge commencing lar furrows, concave border, short palpebral lobes, near middle of anterior glabellar lobe, with fine punctate ornament, and lack of a preglabellar dense punctae weakly evident on some specimens. field. Posterior border of fixigenae narrow (exsag.) Occurrence.—Prodamesella is known only and convex; posterior border furrow long and from the Mapania Zone in central Shantung, shallow with slight anterolatcral curve. China, and from the Peronopsis opimus Zone in Remarks.—This species is distinguished from northwestern Queensland. P. convexa Chang by its shorter anterior border, medially indented and slightly expanded anterior PRODAMESELLA BISERRATA Jell, n. sp. glabella, more oblique IP and 2P furrows, and Plate 1, figure 9; Plate 4, figures 1-6 less prominent genal ornament. Holotype.—Cranidimu UQF69363 from UQL Damage to the occipital area of illustrated 256 (Pl. 4, fig. 2). cranidia of P. convexa and lack of knowledge of Other material.—Eleven cranidia, including other exoskcletal parts for either species limits the figured specimens UQF3196, 69362-69367, are understanding of the species as well as the genus. known from the type locality. One other speci- Four pits surrounding a small tubercle is an men is known from Queensland Museum locality occipital feature apparently unknown in other 122 (lat. 19°32' S., long. 138°54' E.) and was Cambrian trilobites except rare olcnids (Palmer, listed as "Ptychopariidae sp." by Jell (1975, p. 6). 1968, p. B68); however, it is more common in Diagnosis.—Prodamesella with short anterior later trilobites, especially the Odontopleuridae border; indented (bibbed) anterior glabella, three and Scutelluidae. Its function is unknown. pairs of prominent lateral glabellar furrows, long Family NEPEIDAE Wh itehouse, 1939 occipital ring with four posteromedial pits, and narrow and weakly punctate fixigenae. Genus PENAROSA Opik, 1970 Description.—Cranidium of small size (maxi- PENAROSA RETIFERA bpik mum length 2 mm), length to width ratio 2:3; Plate 4, figures 17-19 moderately convex with steep slope near border Penurosa retilera ôpiK, 1970, p. 25-29, pl. 8, figs. 1, 2; furrow; axial furrow deep laterally, but shallower pl. 9, figs. 1-4; pl. 17, figs. 7-9.

18 The University of Kansas Paleontological Contributions-Paper 90

Material.-Three badly damaged specimens Description.-The cranidium is subquadrate including a cranidium, fixigena, and part of a and quite convex. Glabella broad (0.43 cranidial thorax. width through palpebral area), subrectangular Discussion.-This material is assigned to Pena- with weak lateral glabellar furrows; reaching an- rosa because of its characteristic long frontal area, terior border furrow. Occipital furrow well im- central boss, upturned border, tubercular orna- pressed, but occipital ring broken. Anterior mentation, eye ridges, and other features. It be- border short, convex, of uniform width (sag. and longs to Opik's (1970, p. 45) informal group exsag.). Fixigenae convex, of moderate width, 4(b) because of its rimless border, and is tenta- tapering slightly forward, crossed by distinct tively assigned to P. retif era rather than P. vittata eye ridge running obliquely to anterior palpebral of that group because of its wider (exsag.) flap lobe opposite midlength of glabella. Palpebral on the posterior area of its fixigenae, narrower lobe short but quite wide (0.3 fixigenal width), (tr.) palpebral area of its fixigenae, and lack of convex, well defined by palpebral furrow. Orna- a ridge on the frontal part of the preglabellar mentation granulose. field. Remarks.-This specimen is almost certainly immature and generic and specific assignment is Family DORYPYGIDAE? Kobayashi, 1935 unwarranted at this time. It is questionably as- Gen. and sp. undet. signed to the Dorypygidae because of its large Plate 4, figure 7 glabella reaching to the anterior border furrow, Material.-One cranidium UQF69368 from convex border, deep axial furrow, and granulose UQL256. ornamentation.

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Jell & Robison-Revision of a Trilobite Faunule 19

einer Monographic der b6htnischen Trilobiten: K. Lu Ycn-hao, 1960, Kembriyskie otlozheniya Kitaya: Sel. Btihmischen Ges. Wiss. (Prague), Abh., v. 5, 176 p., Rec., n. ser., v. 4, p. 199-216. [Cambrian deposits 7 pl. of China.] Hill, Dorothy, Playford, Geoffrey, & Woods, J. T. (eds.), , Chang Wen-tang, Chu Chao-ling, Chien Yi-yuan, 1971, Cambrian fossils of Queensland: Queensland & Hsiang Lee-wen, 1965, Zhonggo Gemenlei Huashi: Palaeontogr. Soc., 32 p., 15 pl. Zhonggitod Sanyechong: v. 1, p. 1-362, pl. 1-66; v. Howell, 13. F., 1959, Agnostina: in Treatise on inverte- 2, p. 363-766, pl. 67-135, Kexuc Chubanshc [Science brate paleontology, R. C. Moore (ed.), Part 0, Arthro- Publication Co.] (Peking). [Fossils of each group of poda 1, P. 0172-0186, Geol. Soc. Am. and Univ. China: Chinese trilobites.] Kansas Press (New York; Lawrence). Opik, A. A., 1967, The Mindyallan fauna of north- lyshin, N. K., 1953, Srednekembriiskie trilobity Kazakh- western Queen.dand: Bur. Miner. Rcsour. Aust., Bull. stana; chast I: Akatl. Nauk Kazakhstan S.S.R., Inst. 74, v. 1, 404 p.; v. 2, 165 p., 67 pl. Geol. Nauk (Alma-Ata), 226 p., 11 pl. [Middle , 1970, Nepeid trilobites of the Middle Cambrian Cambrian trilobites from Kazakhstan; Part I.] of northern Australia: Bur. Miner. Resour. Aust., Bull. Jago, J. B., 1976, Late Middle Cambrian agnostid trilo- 113,48 p., 171)1. bites from north-western Tasmania: Palaeontology, Palmer, A. R., 1955, The faunas of the Riley Formation v. 19, p. 133-172, pl. 21-26. in central Texas: J. Paleontol., v. 28, p. 709-786, pl. Jell, P. A., 1975, Australian Middle Cambrian codiscoids 76-92. [Dated November 1954; mailed January 15, with a review of the superfamily: Palaeontographica, 1955.] Abt. A, v. 150, 97 p., 29 pl. , 1968, Cambrian trilobites of east-central Alaska: Khalfin, I.. L. (cd.), 1955, Atlas rukovodyashchikh form U.S. Geol. Surv., Prof. Pap. 559-B, p. 1-115, pl. 1-15. iskopaernykh fauny i flory zapadnoy sibiri: Gosgeol- , & Gatehouse, C. G., 1972, Early and Middle Cam- texizdat, Moskva, 502 p., 85 pl. [Atlas of leading brian trilobites from Antarctica: U.S. Geol. Sun., Prof. fossil fauna and flora of western Siberia.] Pap. 456-D, p. 1-37, pl. 1-6. , 1960, Biostratigrafiya Paleozoya Sayano-Altayskoy Pck, Ilja, & Vanek, Jiff, 1971, Revision of the genera gornoy oblasti; tom 1, Nizhniy Paleozoy: Sibirskiy Peronapsis Hawk et Corda, 1847, and Diplorrhina Nauchno-Issled. Inst. Geol. Geofiz. Miner. Syrya Hawle et Corda, 1847 (Trilobita) from the Middle (SNIIGGIMS), Trudy, v. 19, p. 1-498, pl. 152-253. Cambrian of Bohemia: Wstn. estfed. Ost. Geol., v. [Biostratigraphy of the Paleozoic of the Sayan-Altai 46, p. 269-276, pl. 1, 2. mountain region; Volume I, Lower Paleozoic.] Paulsen, Christian, 1927, The Cambrian, Ozarkian and Kobayashi, Teiichi, 1935, The Cambro-Ordovician forma- Canadian faunas of northwest Greenland: Mcdd. om tions and faunas of South Chosen. Palaeontology. Gronland, V. 70, no. 2, p. 233-343, pl. 14-21. Part Ill: J. Fac. Sci. Tokyo Univ., sec. 2, v. 4, p. 49- 1960, Fossils from the late Middle Cambrian 344, pl. 1-24. Bolaspidella Zone of Mendoza, Argentina: Mat.-fys. , 1939, On the agnostids (Part 1): J. Fac. Sci. Medd. dan. Vidensk. Selsk., v. 32, p. 1-42, pl. 1-3. Tokyo Univ., sec. 2, v. 5, p. 69-198. Rasetti, Franco, 1948, Middle Cambrian trilobites from , 1960, The Cambro-Ordovician formations and the conglomerates of Quebec: J. Palcontol., v. 22, p. faunas of South Korea, Part VII, Palaeontology VI: 315-339, pl. 45-52. J. Fac. Sci. Tokyo Univ., sec. 2, v. 12, p. 329-420, pl. 19-21. Raymond, P. E., 1913, On the genera of the Eodiscidae: , 1962, The Cambro-Ordovician formations and Ottawa Nat., v. 27, p. 101-106. faunas of South Korea, Part IX, Palaeontology VIII: Repina, L. N., Petrunina, Z. E., & Hajrullina, T. I., 1975, J. Fac. Sci. Tokyo Univ., sec. 2, v. 14, p. 1-152, pl. 1-8. Trilobity: Inst. Geol. Geofiz. Sibirskiy Otd. (IGIG), Lazarenko, N. P., 1965, Nekotorye novye srednckem- Trudy, v. 278, P. 100-248, pl. 7-48. [Trilobites, in briyskie trilobity severa sredney sibiri: Nauchno- Stratigraphy and fauna of lower Paleozoic, the north- Issled. Inst. Geol. Arktiki (NIIGA), Uch. Zap., v. 7, ern submontane belt of Turkestan and Alai ridges.] p. 14-36, pl. 1-3. [Some new Middle Cambrian Resser, C. E., 1938, Cambrian System (restricted) of the trilobites from the north of Central Siberia.] southern Appalachians: Geol. Soc. Am., Spec. Pap. 15, , & Datscnko, V. A., 1967, Biostratigrafia verkh- 140 p., 16 pl. nego kembriya scvero-zapada Sibirskoy platformy: Richter, Rudolf, & Richter, Emma, 1940, Studicn in' Nauchno-Issled. Inst. Geol. Arktiki (NIIGA), Uch. Paliiozoikum der Mittelmeer-Linder, 5, Die Sattkianda- Zap., v. 20, p. 13-32. [Upper Cambrian biostratigra- Stufe von Andalusien, eine frcmde Fauna im curo- phy of the northwest Siberian platform.] piiischen Ober-Kambrium: Senckenb. naturforsch. Gcs., , 8c Nikiforov, N. I., 1968, Kompleksy trilobitov ix Abh. 450, p. 1-81, pl. 1-5. otlozheniy verkhnego kembriya reki Kulyumbe (severo- Robison, R. A., 1964, Late Middle Caniltrian faunas from zapad Sibirskoy platformy): Nauchno-Issled. Inst. western Utah: J. Palcontol., v. 38, p. 510-566, pl. Geol. Arktiki (NIIGA), Uch. Zap., v. 23, p. 20-71, 79-92. pl. 1-15. [Trilobite assemblages from Upper Cambrian , 1976, Middle Cambrian biostratigraphy of the deposits of the Kulyumbe River region (northwestern Great Basin: in Paleontology and depositional environ- Siberian plattorn:).] ments, Cambrian of western North America, R. A. Lachman, Christina, & Duncan, Donald, 1944, Early Robison & A. J. Rowell (eds.), Brigham Young Univ.

Upper Cambrian faunas of central Montana: Geol. Geol. Studies, y. 23, pt. 2, p. 93 - 109. Soc. Am. Spec. Pap. 54, 181 p., 19 pl. , 1978, Origin, taxonomy, and homeomorphs of

20 The University of Kansas Paleontological Contributions—Paper 90

Doryagnostus (Cambrian Trilobita): Univ. Kansas eastern Australia, Parts 1 and 2: Queensland Mus., Paleontol. Contrib., Pap. 91, p. 1-10, pl. 1-2. Mem., v. 11, p. 59-112, pl. 8-10. Rozova, A. V., 1964, Biostratigrafiya i opisanie trilobitov , 1939, The Cambrian faunas of north-eastern .o.ednego i verkhnego kembriya severo-zapada Sibirskoy Australia, Part 3: Queensland Mus., Mem., (n. ser.) platformy: Inst. Geol. Geofiz. Sibirsk Otd. Akad. Nauk v. 1, p. 179-282, pl. 19-25. SSSR (Moscow), 148 p., 19 pl. [Biostratigraphy and , 1945, The Cambrian faunas of north-eastern description of trilobites of the Middle and Upper Cam- Australia, Part 5: Queensland Mus., Mcm., v. 12, p. brian of the north-west Siberian platform.] 117-123, pl. 11. Shergold, J. H., Druce, E. C., Radke, B. M., & Draper, J. J., 1976, Cambrian and Ordovician stratigrapby of Peter A. Jell the eastern portion of the Georgina basin, Queensland Department of Geology and eastern Northern Territory: 25th Int. Geol. Congr., University of Queensland Field Excursion Guide!). 4C, 54 p. St. Lucia, Queensland 4067 Westergard, A. H., 1946, Agnostidea of the Middle Cam- Australia brian of Sweden: Sver. Geol. Unders., ser. C, no. 477, 140 p., 16 pl. Richard A. Robison , 1949, Opsidiscus, a new name replacing Aulaco- Department of Geology and discus Westergtird, 1946: Geol. FOren. Stockholm Paleontological Institute FOrh., v. 71, p. 606. University of Kansas Whitehouse, F. W., 1936, The Cambrian faunas of north- Lawrence, Kansas 66045

EXPLANATION OF PLATES Unless indicated otherwise, specimens on all plates are from locality UQL256 of the Currant Bush Limestone (late Middle Cambrian), northwestern Queensland.

PLATE 1 FIGURE latex cast of holotype, partially exfoliated (figured FIGURE by Whitehouse, 1936, pl. 8, fig. 11), X6, 1,2,4,8. Baltagnostus damesi (Rcsser & Endo), Mapan UQF3195.-5. Small holaspid pygidium, X8, Formation, Middle Cambrian, Manchuria (all X8 UQF69637. and all numbered USNM 57829.-1a,b. Dorsal 7-9,12. Doryagnostus deltoides Robison, n. sp.-7a,b. and lateral views of cephalon with incomplete Dorsal and lateral views of latex cast of cephalon preglabellar median furrow (cf. with retouched (figured as paratype of Euagnostus opinuts by illustration by Endo & Resser, 1937, pl. 30, fig. Whitehouse, 1936, pl. 8, fig. 10), X6, UQF3194. 1).-2a,b. Stereogratn of cephalon with incom- —8a-c. Lateral view and stereogram of partly plete preglabellar median furrow (cf. with re- exfoliated pygidium, X7, UQF69625.-9a,b. touched illustration by Endo & Resser, 1937, pl. Stereogram of holotypc cephalon, X6, UQF69624. 30, fig. 2).-4a-c. Lateral view (a) and stereo- —12. Pygidium, X5, UQF69626. gram (b, c) of lectotype pygidium (figured by 10. Hypagnostus? sp. Damaged and partly exfoliated Endo & Resser, 1937, pl. 30, fig. 4).-8. Pygidium cephalon, x9, UQF69638. with broken border spines (figured by Endo & 11. Hypagnostus cf. H. &pens Whitehouse, V Creek Resser, 1937, pl. 30, fig. 3). Limestone, Middle Cambrian, Queensland. Pygid- 3,5-7, Baltagnostus attstralis Robison, n. sp., all X8.— ium, X6.5, UQF69639. 9-11. 3,11. Representative cephala, UQF69627, 69628. —5a-c. Lateral view (a) and stercogram (b, c) PLATE 3 of holotypc cephalon, UQF69629.-6a-c. Lateral view (a) and stereogram (b, c) of slightly ex- Chondranomocare conf es-turn (Whitehouse) foliated pygidium, UQF69630.-7,10. Medium FIGURE and small holaspid pygidia, UQF69631, 69632.— /. Left pleura and part of axis of damaged thoracic 9a,b. Stereogram of pygidium (figured as para- segment, X8, UQF69350. type of Euagnostus opinuts by Whitehouse, 1936, 2-5. Growth series of small cranidia showing develop- pl. 8, fig. 12) associated with cranidium of ment and lengthening of preglabellar field and Prodamesella biserrata, n. sp., UQF3196. decrease in relative size of the eyes; X24, X15, X6, and X6, UQF69351-69354, respectively. Fig- PLATE 2 ure 3 is a damaged complete specimen from UQL463, Currant Bush Limestone. FIGURE 1-6. Pcrono psis opimus (Whitehouse).—la-c. Stereo- 6. Posterior oblique view of posterior pygidial frag- gram (a, b) and lateral view (c) of well-preserved ment (figured by Whitehouse, 1939, pl. 23, fig. cephalon, X6, UQF69633.-2,3,6. Holaspid ceph- 25), X5, UQF3357. ala of various sizes, all X6, UQF69634-69636.— 7,8,12. Pygidia showing posterolateral angularity, wide 4a-c. Stereogram (a, b) and lateral view (c) of doublure, steep posterior slope, and postaxial keel.

Jell & Robison—Revision of a Trilobite Faunule 21

FIGURE PLATE 4 —7b. Left lateral view.-8b. Posterior oblique FIGURE view. X2.5, X6, X5; UQF69355-69357, re- 1-6. Prodamesella biserrata Jell, n. sp. Cranidia; all spectively. X12, except 2b, which is X10; UQF69362-69367, 9. Latex cast of damaged pygidium (figured by respectively.-2a,b. Dorsal and anterolateral ob- Whitehouse, 1939, pl. 23, fig. 27), X6, lique views of holotypc (UQF69363).-1,3-6. UQF3359. Paratypes. 10. Damaged pygidium (figured by Whitehouse, 1939, pl. 23, fig. 26), X5, UQF3358. 7. Gen. and sp. undet. Cranidium, X12, UQF69368. 11. Holotype cranidium (figured by Whitehouse, 1939, 8. Opsidiscus microspinus Jell. Cranidium, X22, pl. 23, fig. 22), UQF3354; a, posterior view UQF69369; a, dorsal, and b, antcrolateral oblique showing posterior fixigenae well below palpcbral views. lobes, X2.5; b, anterolateral oblique view, X3; 9-16. Sudanomocarina changi Jell, n. gen. & sp.— c, dorsal view, X3. 9,10,12,14,16. X8, X4 (10a, X5), X4, X6, 13. Cranidium, X3, UQF69358; a, oblique lateral UQF69370, 69371, 69373, 69375, 69377, view; b, dorsal view. X6; respectively.-11. Pygidium, X7, UQF69372.— 14. Cranidium (figured by Whitehouse, 1939, pl. 23, 13,15. Damaged juvenile cranidia, X 8 and X10, fig. 23), X3, UQF3355. UQF69374 and 69376, respectively. Figure 14 15. Hypostoma, X6, UQF69359. (UQF69375) is the holotype. Specimens in 9,10, 16. Librigena, X3, UQF69360. 12,14,16 are from UQL463, Currant Bush Lime- 17. Cranidium (figured by Whitehouse, 1939, pl. 23, stone. fig. 24), X3, UQF3356; a, latex cast; b, damaged original. 17-19. Penarosa rctifera Dpik.-17. Damaged cranidium, 18. Latex cast of cranidium, X3, UQF69361. X4, UQF69378.-18. Fragment of right fixigena, 19. Latex cast of librigena (figured by Whitehouse, X5, UQF69379.-19. Thoracic fragment, X5, 1939, pl. 23, fig. 28), X4, UQF3360. UQF69380. THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS Paper 90, Plate 1 Jell & Robison Revision of a Trilobite Faunule

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8b 8c 12 THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS Paper 90, Plate 3 Jell & Robison--Revision of a Trilobite Faunule THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS Jell & Robison—Revision of a Trilobite Founule Paper 90, Plate 4