Lower and Middle Cambrian trilobites from the Digermul peninsula, Finnmark, northern Norway

FRANK NIKOLAISEN & GUNNAR HENNINGSMOEN

Nikolaisen, F. & Henningsmoen, G. 1987: Lower and Middle Cambrian trilobites from the Oiger­ mul peninsula, Finnmark, northern Norway. Nor. geol. unaers. Bul/. 419, 55-95.

Ten trilobite species are described. The only Lower Cambrian one is the olenellid Kjerulfia lata from the upper member (02) of the Ooulbasgaissa Formation, previously known from southern Nor­ way. The Middle Cambrian species are from the two lower members (K1, K2) of the overlying Kistedal Formation. The trilobites from Kl are described as Eflipsocephalus cf. tiottlt. Eccaparadoxides cf. pusillus and Hydrocephalus cf. carens and are compared to Czechoslovakian and Polish species. The trilobites from K2 are Doryagnostus tncertus, first described from sout­ hern Norway, the subspecies Peronopsis terox sal/esi , earlier known from France and Spain, Paradoxides davidis, earlier known from various countries including southern Scandinavia, a new species, Nassovia? mjol/nir n.sp., and two incompletely known forms, described below as Ptychopa­ riinae gen. et sp. indel. and Anomocarina? sp.. Thus, both Middle Cambrian trilobite faunas differ from the very well documented contemporaneous ones in southern Scandinavia, but especially the fauna from K1. A striking difference is also the scarcity of agnostid trilobites.

Frank Nikolaisen & Gunnar Henningsmoen, Paleontologisk museum, Sars gate " N-0562 Oslo 5, Norway.

Introduction University carried out fieldwork financed by Upper Cambrian and lower Tremadoc trilobites the Geological Survey of Norway. Further fos­ (all olenids) from the Digermul peninsula were sils were collected on expeditions from Oxford described in an earlier paper (Nikolaisen & University in 1961 and 1963 (Reading 1965, Henningsmoen 1985). The present paper on pp. 168, 170) and subsequently presented to Lower and Middle Cambrian trilobites conclu­ the Palaeontological Museum, University of des the description of the hitherto known trilobi­ Oslo. The present authors made their own tes from the peninsula. collections in 1960 (Henningsmoen 1961) and The Digermul peninsula (Digermulhalv0ya) again in 1963 when they were accompanied is situated in the Tana area, Finnmark county, by Amanuensis Kari E. Henningsmoen. Field­ northern Norway (Fig. 1). It is 30 km long and work was financed by the Fridtjof Nansen separates Langfjord from the inner part of Fond and associated funds. Tanafjord. In 1980 and later, collections have been An overall presentation of the geology in the made by B.-D. Erdtmann and others from the Tana area was given by S. F0yn (1937). H.G. Georg-August-Universitat, Gottingen, western Reading (1965) gave a detailed account of the Germany, and the trilobites from these have geology of the Digermul peninsula and descri­ been presented to the Palaeontological Muse­ bed the stratigraphy of the Eocambrian (Vendi­ um in Oslo. an),Cambrian and Tremadoc rocks exposed. The present study of the Lower and Middle Fossils were first found on the peninsula in Cambrian trilobites was initiated by the senior 1934 by Sven F0yn (Feyn 1937) - brachio­ author (G.H.) in 1965, but the work was inter­ pods, a hyolithid and a trail of Cruziana type, rupted until 1984 when the junior author (F.N.) described by Trygve Strand (1935) and assig­ took over the task to complete and extend the ned to the Middle Cambrian. The first trilobite, description, with some participation by the a paradoxidid, was collected by Harold G. senior author. Reading in 1950 (Reading 1965, p. 168). Up­ per Cambrian and lower Tremadoc olenid trilo­ bites and lower Tremadoc graptolites were Techniques discovered in 1959 by R. Pattinson and J.K. Where necessary, specimens have been pre­ Russell, who with H.G. Reading from Oxford pared using a vibro drill or needles. Linear 27"50 28'00' 28'10' 28'20

27"50' 28'00' 28'10' 28"20 Fig. 1. Sketch map of Digermul peninsula (Digermulhalv"ya), Tanafjord, Finnmark, northern Norway, with main samp­ ling sites (1-17) for Lower and Middle Cambrian trilobites. Inset map shows location of the Digermul peninsula (ar­ row). Concerning geographical names in Finnmark, both Lappish and Norwegian names are official, but only the latter are used here. 1. Lower Cambrian. Greenish-grey micaceous siltstone with flakes of greenish shale interbedded in the middle part of the massive bedded quartzite member (02) of the Doulbasgaissa Formation. Scree at outcrops on the northwestern slope of Bardeviktind. 2-4. Lower Middle Cambrian. Quartzite and shale member (Kl) of the Kistedal Formation. 2. Grey or green siltstones interbedded with thin quartzite beds. Eastern slope of Hansvikdat,1 km northwest of Nova. 3. Sediments as in locality 2. Section in creek in northern part of Hansvlxdal, about 1 km south of the sea-shore. 4. Alternating bands of grey quartzite and sandstone with greenish-grey shales, approximately 3 m above the top white quartzite of the Doulbasgaissa Formation. Southeastern slope of Kistedalen. 600 m east of small 'sausage­ shaped' pond. 5-17. Middle and upper Middle Cambrian. Quartzite and shale member (K2) of the Kistedal Formation. 5. Thin-bedded sandstone and grey/greenish-grey shale. About 1 km east of the small lakes Bokselvvannene. 6. Sediments as in locality 5. Creek section 5 km south-southwest of Nova. 7. Sediments as in locality 5. Loose specimens and specimens collected in various horizons from about 25 m to about 50 m abovethe quartzite and shale member (K1).Section in a small stream flowing down into Hansvikda',4 km south of Nova. 8. Sediments as in locality 5, horizons as in locality 7. Southeastern hillside about 800 m northeast of locality 7. 9. Sedimentsas locality 5, horizon as in locality 7. The most southeasterly tributary to H~nsvikdal, 2 km south of l2lrene. 10. Sediments as in locality 5, horizons as in locality 7. The most southerly tributary of Kistedal, nearly 6 km south­ southwest of the summit of Perletind (Berlogaissa). 11. Sediments as in locality 5, probably horizons as in locality 7. Loose specimens from the hillside 3 km northwest of locality 10. 12. Sediments as in locality 5, horizons as in locality 7. Eastern slope of Kistedalen, about 700 m southeast of small 'sausage-shaped' pond. 13. Sediments as in locality 5. Various horizons from about 55 m to about 75 m above the top of the quartzite and shale member (Kl). Western slope of Hansvikdat, 2 km south-southeast of Nova. 14. Sediments as in locality 5, horizons as in locality 13. Southwest of Kistedal, about 4 km southeast of l2lrene. 15. Yellowish shale with articulate brachiopods in abundance. Horizon probably as in locality 13. Loose specimens at northern tributary to Manndraperelva, 5 km south-southwest of the summit of Perletind (Berlogaissa). 16. Darker grey, shaly mudstones. Possibly somewhat younger than the horizons in locality 13. Section along the river in Kisledalen, 1 km northeast of the largest lake in the valley. 17. Darker micaceous mudstone. Probably high up in the quartzite and shale member. Scree along the eastern shore of Kistedal. NGU-BULL.419,1990 Lowerand Middle Cambrian trilobites 57 measurements have been made using either hing their assignment to taxa of highest equal a micrometer ocular or vernier calipers. Angles rank within the Trilobita. were measured using a protractor and are given to the nearest 5°. Order Agnostida Kobayashi, 1935 All specimens have been coated with ammo­ Suborder Agnostina Salter, 1864 nium chloride before photographing. A fluores­ Superfamily Agnostacea M'Coy, 1849 cent ring-light illuminator and an additional Family Agnostidae M'Coy, 1849 northwest source were utilised for illumination. Subfamily Quadragnostinae Howell, The photographs are not retouched. The dra­ 1935(a) wings in this and the 1985 paper were made by the junior author. Genus Peronopsis Hawle & Corda, 1847

Synonyms. - Mesospheniscus Hawle & Cor­ da, 1847 (type species by monotypy: Battus Systematic descriptions cuneifer Barrande, 1846), Diplorrhina Hawle & Terminology Corda, 1847 (type species by subsequent The terminology mainly follows that of Harring­ designation by Snajdr 1958: D. triplicata Haw­ ton et al. (1959, pp. 0117-0126). For brevity la & Corda, 1847 = subjective junior synonym the symbols 'L1·L4' are used for the lateral of Battus cuneifer Barrande, 1846), Mesagnos­ glabellar lobes and 'S1-S4' for the furrows, tus Jaekel, 1909 (type species by original as suggested by Jaanusson (1956, p. 37; also designation: Battus integer Beyrich, 1845), Henningsmoen 1957, p. 12). Other terms appli­ Pseudoperonopsis Harrington, 1938 (type spe­ ed are those given by Henningsmoen (1957, cies by original designation: Agnostus Sal/esi pp. 12-14; 1959, pp. 154-157) and c>pik (1967, Bergeron, 1889), ?Acadagnostus Kobayashi, pp. 52-62), but following the recommendations 1939a (type species by original designation: given by Whittington & Kelly (1983a, pp. 1-29; Agnostus acadicus Dawson, 1868). 1983b, pp. 1-9, Pis. 1-7). However, for the longitudinal furrow dividing the anterior and Type species. - Battus integer Beyrich, 1845. posterior palpebro-ocular ridges we apply the term 'epipalpebral furrow' introduced by Co­ Peronopsis ferox (Tullberg, 1880) sal/esi wie & McNamara (1978, p. 616; also McNama­ (Bergeron, 1889). ra 1978, p. 636) in preference to 'ocular fur­ row' introduced by Sdzuy (1978, p. 93) and Fig.6:A-G to 'palpebral ledge' as suggested for a corre­ sponding deep and strong groove in Lophosa­ 1888 Agnostus Sal/esi n. sp. - Munier-Chal­ uka by Shergold (1972, p. 15). We also ac­ mas & Bergeron, p. 376 (nomen nudum). cept the term 'metagenal ridge' extending from 1888 Agnostus Sal/esi Munier-Chalmas et the axial furrow to the metagenal spine in ole­ Bergeron - Bergeron, p. 284 (nomen nellids as introduced by Bergstrom (1973b, nudum). Fig. 1; also 1973c, p. 284). Finally, we employ 1889 Agnostus Sal/esi Mun.-Chalm. et J. Berg. the term 'deltoid area' introduced by Robison - Bergeron, pp. 337-338, PI. 3, fig. 5 (1978, p. 5) for the triangular depression for­ (descr. and fig. of dorsal exoskeleton). med by the expansion of the anterior border 1935 Peronopsis sal/esi (Munier-Chalmas and furrow at its junction with the preglabellar Bergeron) - Howell, pp. 226-227, PI. median furrow in agnostids. 22, figs. 17-18 (descr. and figs. of two dorsal exoskeletons). Classification 1935 Peronopsis Sal/esi Munier-Chalmas et The systematic classification mainly follows Bergeron - Thoral, pp. 38-39 (remarks). that of Harrington et al. (in Moore 1959), but 1938 Pseudoperonopsis sal/esi (M. Chalmas with a few modifications according to more et Bergeron) - Harrington, p. 151 (erects recent studies. We thus prefer mainly to follow Pseudoperonopsis with A. Sal/esi as type Oplk's (1967; 1979) classification of agnostids. species). We fully agree with Bergstrom (1973a, p. 37) 1939a Peronopsis sal/esi (Munier-Chalmas and that the olenellids and redlichiids should be Bergeron, 1889) - Kobayashi, p. 115 grouped in separate orders, thus re-establis- (listed). 58 Frank Nikolaisen & Gunnar Henningsmoen NGU·BULL419,l990

1939b'Peronopsis (Pseudoperonopsis) sal/esi Dimensions: - The largest cephalon (PMO (Munier-Chalmas et Bergeron) - Kobay­ 111.589) is 3.4 mm long and 4.0 mm wide, ashi, p. 579 (remarks on the systematic and the largest pygidium (PMO 111.588) is position of Pseudoperonopsis). 4.0 mm long and 4.3 mm wide (twice the width 1958 Peronopsis fal/ax seues: (Munier-Chal­ of right half). mas & Bergeron 1889) - Lotze, pp. 732, 738 (listed). Remarks. - Pygidia in the Finnmark material 1961 Peronopsis teuex sal/esi (Munier-Chal­ agree almost to the finest details with pygidia mas & Bergeron 1889), - Sdzuy, pp. from Montagne Noire, southwestern France, 523-524 (241-242), PI. 2, figs. 1-8, PI. as described and figured by Courtessole (1973, 28, fig. 15 (comparison with P. f. ferox pp. 116-117, PI. 2, figs. 4-6, 8-9, 11, 13) and and figs. of cephala and pygidia). from Los Barrios de Luna, northwestern Spa­ 1961 Peronopsis fal/ax - Henningsmoen, p. in as figured by Sdzuy (1961, PI. 2, figs. 4-8) 94 (recorded from Finnmark). and GiI Cid (1981, PI. 1, fig. 3). The only diffe­ 1965 Agnostids [partim.] - Reading, p. 175 rence that may be observed is that the margi­ (listed from Finnmark). nal spines are more slender in the Finnmark 1967 Peronopsis sal/esi M.Ch. et Berg. ­ material. Courtessole, pp. 492 (2), 515 (23), 519 We agree with Robison (1982, pp. 153-155) (25), 521 (27) (listed). in that several forms hitherto referred to as 1971 Agnostids [partim.] - Henningsmoen, subspecies of P. fal/ax (Linnarsson, 1869), and p. 90 (reported from Finnmark). among them P. fal/ax ferox, should warrant 1973 Peronopsis fal/ax sal/esi (Munier-Chal­ elevation to specific rank. On the other hand, mas y Bergeron 1889) - Courtessole, we do not follow him (op, cit. p. 155) in consi­ pp. 116-117, PI. 2, figs. 4-14, text-fig. dering P. fal/ax ssttes! to be a synonym of 21/5 (descr. and figs. of dorsal exoskele­ P. ferox. The differences between pygidia of tons, cephala and pygidia). P. ferox and those of the French-Spanish form 1974 Agnostids [partim.] - Martinsson, p. 237 seem fairly obvious and quite well defined. (mentioned from Finnmark). However, we do not regard these differences 1981 Peronopsis fallax sal/esi (Munier-Chal­ as reflecting specific, but rather subspecific mas y Bergeron, 1889) - Gil Cid, pp. characteristics. Sdzuy (1961, pp. 523-524) did 118-119, PI. 1, fig. 3 (descr. and fig. of not, as stated by Opik (1979, p. 43) regard pygidium). P. sal/esi as a subspecies of P. ferox but of 1982 Peronopsis fallax ssttesi (Munier Chal­ P. fal/ax. Sdzuy did, however, point out the mas and Bergeron) - Robison, p.155 close similarity and presumed relationship (considered a synonym of P. ferox). between P. seltes! and P. terox. P. ferox ferox 1986 Peronopsis terox (Tullberg, 1880) ­ occurs in dark, fine-grained shales, whereas Unan & Gozalo, pp. 42-43, stratigraphi­ P. f. sal/esi is found in lighter shales em­ cal distribution in sections in Figs. 10 (p. bedded in sandstones or quartzites both in­ 23), 11(p. 25), 12(p. 26), PI.2, figs. 11-14. southwestern Europe and in Finnmark. Thus, it may well be that the two subspecies have Type specimen. - Holotype by monotypy is been confined to two different types of environ­ the dorsal exoskeleton from an uncertain Midd­ ments. le Cambrian zone in Montagne Noire, France, Both Munier-Chalmas & Bergeron's (Janua­ figured by Bergeron 1889, PI. 3, fig. 5, preser­ ry 1888) and Bergeron's (February 1888) publi­ ved at the Laboratoire de geologie de la Sor­ cations were only short reports without de­ bonne, Paris, France. scriptions or illustrations. The specific taxon sal/esi therefore became valid with Bergeron's Finnmark material. - Sixteen more or less (1889) description and illustration of an almost fragmentary cephala, one external mould of a complete dorsal exoskeleton. The latter work pygidium with articulated but poorly preserved was written solely by Bergeron and hence he thorax and thirteen other pygidia, all from the is the only author attributed to the taxon. middle Middle Cambrian shale and sandstone Oplk (1979, p. 43) discussed the validity of the member (K2) of the Kistedal Formation at loca­ use of the name P. seues! according to Ho­ lites 7, 11 and 12. well's (1935, p.226) information on the inaccu­ rate original description and illustration. Opik NGU • BULL. 419. 1990 LowerandMiddle Cambrian trilobites 59 stated, however, on the basis of subsequently Jianshan, Zhejiang, China, figured by Yang described and figured plesiotype material by (1982, PI. 2, fig. 3) as P. taitzuhoensis Lu Howell (1935, pp. 226-227, PI. 22, figs. 17-18) (1957, p. 258, PI. 137, figs. 4-5; also Lu et and Sdzuy (1961, pp. 523-524 [241-242], PI. al. 1965, p. 49, PI. 5, figs. 21-22). Both Chine­ 2, figs. 1-8, PI. 28, fig. 15) that "the name se forms are rather small and reliable differen­ sallesi attributed to it is a reasonable but sub­ ces between them and P. ferox sallesi can jective inference". Nevertheless, Opik maintai­ hardly be given here. The pygidium of P. tait­ ned Harrington's (1938)generic name Pseudo­ zunoensis (which lacks the posterior axial fur­ peronopsis while waiting for the holotype of row as seen in Lu's original) seems, howe­ P. sallesi to be redescribed and refigured. ver, to be slightly shorter than that of the Finn­ Accepting the plesiotype material to be suffici­ mark form. ent for the recognition of the name P. sallesi, we prefer to reject the name Pseudoperonop­ Subfamily Euagnostinae Opik. 1979 sis because material of this fulfils the generic concept of Peronopsis as given by Robison Genus Doryagnostus Kobayashi. 1939(a) (1964, p. 530), but also Opik's own (1979, p. 53), except for his statement that "(11) the Synonym. -Ceratagnostus Whitehouse, 1939 posterior axial node is laterally expanded and (type species by original designation: C. magis­ its flanks are convex". This latter character is ter Whitehouse, 1939). fairly vague even in the type species and is also only very faintly present in some speci­ Type species. - Agnostus incertus Bmgger, mens in the Finnmark material (Fig. 6:C) and 1878 by original designation. in Sdzuy's material (1961, PI. 2, fig. 6). Furthermore, Oplk (1979, pp. 43-44) empha­ Doryagnostus incertus (Bmgger, 1878) sized the similarity between Linguagnostus perplexus Robison, 1964 (transferred to Pseu­ Figs. 2a-c, 6:H-M doperonopsis by Opik) from the Bolaspidella contracta Subzone in western Utah and V­ 1878 Agnostus incertus, n. sp. - Br0gger, Creek Limestone, Quita Formation of Queens­ pp. 70-71, PI. 6, figs. 4a-b (descr. and land, Australia, and P. f. sa/lesi. L. perplexus figs. of cephalon and pygidium). differs, however, from the latter in having a 1880 Agnostus incertus Bmgger - Tullberg, wider pygidial axis, and a slightly shorter p. 19, PI. 1, figs. 6a-b (descr. and figs. (sag.) postaxial field and a better defined ante­ of cephalon and pygidium). rior pygidal segment. 1883 Agnostus incertus Bragger - Linnars­ P. bifidus Khajrullina (in Repina et al. 1975, son, p. 32 (recorded). pp. 109-110, PI. 9, figs. 12-15, PI. 10, figs. 1890 Agnostus incertus Bragg. - Holm, p. 1-3) from the upper Middle Cambrian Amginsk 267 [13] (listed). and Majsk Formations in the northern sub­ 1902 Agnostus incertus Br. - Granwall, pp. montane belt of Turkestan is rather similar to 52-53 (recorded). P. ferox sallesi. Their cephala can hardly be 1907 Agnostus incertus, Bragger - Lake, pp. separated, but pygidia of P. f. sallesi differ 29-30, PI. 3, figs. 1-3 (descr. and figs. from those of the Turkestan species in having of cephala and pygidium). a wider axis and narrower (sag.) postaxial fi­ non 1916 Agnostus et. incertus Bragger - 11­ eld and in having the posterior border far less ling, p. 407, PI. 28, fig. 10 (remarks and distinctly zoned (see definition by Opik 1967, fig. of pygidium ) [= Peronopsis scutalis p. 69). scutalis (Hicks, 1872)according to Rush­ Strikingly similar to the present form are ton 1979, p.50]. both the holotype of P. majiangensis Lu & 1929 Agnostus incertus Br. - Strand, pp. Chien (in Lu et al. 1974, p. 100, PI. 39, fig. 2; 344-345 (remarks). also Lu & Chien in Yin & Li 978, p. 390, PI. 1930 Agnostus incertus Bragg. - WaJlerius, 144, fig. 27) and additional conspecific materi­ pp. 52, ?54 (recorded). al (Lu & Qian 1983, pp. 21-22, PI. 1, figs. 1935 Agnostus incertus Bragger - Thorslund, 13-14, PI. 2, fig. 9) from the southeast Guiz­ p. 107, PI. 1, figs. 9-10 (recorded and nou southwestern China, and the pygidium figs. of cephalon and pygidium). from the Yangliugang Formation at Duibian in 60 Frank Nikotsiset: & Gunnar Henningsmoen NGU • BULL. 419. 1990

1939a Doryagnostus incertus (Bragger) - Ko­ red) and do not show any details that could bayashi, p. 148 (erection of Doryagnos­ help to identify the specimens, and no catalo­ tus). gue numbers were given. 1946 Doryagnostus incertus (Bragger, 1878) Westergard (1946, p. 83-84) remarked that - westercaro, pp. 83-84, PI. 12, figs. the pygidial axis varies in breadth in his speci­ 20-23, PI. 13, figs. 1-3 (remarks and figs. mens of D. incertus from Sweden, but that of cephala and pygidia). none of them had an axis as broad as in 1948 Doryagnostus incertus (Bmgger) ­ Bmgger's figure. The same is, however, true Strand, p. 92 (recorded). of the specimens from Krekling in our collecti­ 1960 Doryagnostus incertus (Bragger) - Po­ ons; the axial width may have been exaggera­ krovskaya (in Chernysheva), PI. 1, figs. ted in Bmgger's drawing. 15-16 (copies of Westergard 1946, PI. A lectotype (PMO 28200, Fig. 2a) and two 12, figs. 22-23). paralectotypes (PMO H2646, Fig. 2b and PMO 1961 Doryagnostus incertus - Henningsmo­ H2646, Fig. 2c) are selected here among speci­ en, p. 94 (recorded from Finnmark). mens collected by Bmgger at Krekling in 1877. non 1962 Doryagnostus incertus (Broegger) ­ They are accompanied by a label with Br0g­ Hutchinson, p. 87, PI. 10, figs. 9-11 (re­ ger's own handwriting. marks and figs. of cephalon and pygi­ Robison (1978) quoted a statement from the dial [= D. magister (Whitehouse, 1939) Paleontological Museum in Oslo that represen­ according to Opik 1979]. tatives of Agnostus incertus collected by Br0g­ 1965 Agnostids [partim.] Reading, p. 175 (li­ ger were missing in its collections. Luckily, sted from Finnmark). quite a number of specimens have now been 1971 Doryagnostus incertus - Henningsmo­ traced, - partly labelled as such by Breqqer en, p. 90 (reported from Finnmark). and partly (PMO H2646) on a piece of shale 1974 Agnostids [partim.] - Martinsson, p. 237 labelled Paradoxides rugulosus by Br0gger (listed from Finnmark). (with the cranidium figured by Bmgger 1878, non 1977 Doryagnostus incertus (Bragger) ­ PI. 2, fig. 1). Other specimens do not have Zhou et al., p. 111, PI. 36, figs. 20-21 labels written by Br0gger, but it is stated that (short descr. and figs. of cephalon and they were collected by Bmgger (alone or with pygidium)[? =D.notalibrae Opik, 1979]. eo-collectors). Further specimens have been non 1978 Doryagnostus incertus (Bragger) ­ added to the collections subsequently. Robison, p. 7, PI.2, figs. 1-2, 8 (descr. and figs. of dorsal exoskeletons) [=0. Finnmark material. - Sixteen more or less magister (Whitehouse, 1939)] complete cephala, thirteen more or less frag­ non 1978 Doryagnostus incertus (Bragger) ­ mentary or distorted pygidia, and one pygidium Yang, pp. 20-21, PI. 1, figs. 11-12 (short articulated with but very fragmentary thorax descr. and copies of figs. of Zhou et al.) and posterior part of the cephalon, all preser­ [? = D. notalibrae Opik, 1979]. ved in greenish shale of the Middle Cambrian 1979 Doryagnostus incertus (Bragger, 1878) shale and sandstone member (K2) of the Kiste­ - Opik, p. 82 (remarks and comparison dal Formation at localities 7, 11, 14, 15 and 18. with D. magister Whitehouse, 1939). 1984 Doryagnostus - Henningsmoen, pp. Dimensions. - The largest cephalon (PMO 23-24 (mentioned and fig. of cephalon 86437) is 4.8 mm long and 5.2 mm wide (twi­ from Finnmark). ce the width of the right half). The largest 1985 Doryagnostus incertus - Berg-Madsen, pygidium (PMO 111.605) is 5.8 mm long and p. 359 (listed). 6.0 mm wide.

Type material. - Bmgger (1878) based his Remarks. - The Finnmark material agrees description of the species on isolated cephala very well with the material from southern Nor­ and pygidia in black shale from Krekling, Ei­ way housed In the PaleontoJogisk Museum in ker-Sandsvcer area, Oslo Region. He thus did Oslo and that from Scania, Sweden, as descri­ not base his description on a single speci­ bed and figured by Westergard (1946, pp. men, but did give drawings of a cephalon and 83-84, PI. 12, figs. 20-23, PI. 1-3, figs. 13). a cranidium. The drawings appear somewhat Although the distance from the posterior end schematic (and are possibly somewhat resto- of the pygidial axis to the posterior border NGU · BUL L. 419. 1990 Lowerand Middle Cambrian trilobites 61

Fig. 2. Doryagnostus incertus (Bmgger. 1978). Black alum shale. Zone of Ptychagnostus punctuosus , Paradoxides paradoxissimus Stage. middle Middle Cambrian. Railroad section at Krekling Station. southern Norway. a. dorsal view of lectotype. almost complete but flattened pygidium. PMO 28200. Dorsal views of b. paralectotype. incomplete and flattened cephalon, and c. paralectotype. practically complete but flattened pygidium. both occurring on the same piece of shale as the cranidium figured by Bmgger 1878. PI. 2. fig. 1 as Paradox ides rugulosus Corda, 1847 [= Eccaparadoxides brachyrachis (Linnarsson . 1882)]. PMO H2646. All latex casts of external moulds. x 8. furrow is slightly longer in the Finnmark speci­ with the synonymy proposed by Westergard. mens than in those from southern Scand ina­ We have examined severa l spec imens collec­ via, we do not find any reason to regard this ted by Br0gger or Br0gger et al. and subsequ­ rather minor difference as being more than ently labelled as paratypes, and also additio­ natural variation between separated populati­ nal topotypes. By far the greater number of ons, and do not hesitate to include our materi­ the pygidia are dist inctly wider than long, the al in the same spec ies. location of the pygidial marginal spines, which Argumentation for a synonymy of D. incertus are undeflected where preserved , vary from and the Australian species D. magister (White­ an anterior to a posterior pos ition to the trans­ house , 1939, pp. 256-257 , PI. 25, fig. 27) was verse level of the axial tip. Well preserved brought forward by Westergard (1946, pp. topotype cepha la also exhibit a deltoid area. 82-83) and more recent ly by Robison (1978, The spec imens from Finnmark have, as the p. 7) who re-illustrated Whitehouse's type Australian form, a slightly larger average size. specimens together with additional topotypes. The only morphological differences between Opik firstly concurred with (1957, p. 14) but Scandinavian and Austra lian specimens that later challenged (1979, p. 82) WestergArd 's we can find are that the Scandinavian pygidia view . He stated that D. magister differs from have a more strongly curved posterior margin D. incertus in having large, undeflected, cur­ (also the spec imens with the marginal spines ved, pygidia l border spines visibly behind the situated far back ), the axial node situated trans verse level of the axial tip and that its more posteriorly and slightly smaller marginal pygidium is wider than long. Opik also pointed spines. We have not found any specimen, out (ibid.) that D. incertus is confined to the neither among the topotype nor the Finnmark Zone of Ptychagnostus punctuosus in Scandi­ material , with a posterior margin as gently navia, whereas D. magister is bi-zonal in Au­ curved as in those from Austral ia. It therefore stralia. Robison (1978, p. 7), moreover, added may seem justified not to synonymize the two that the Australian specimens differ from tho­ forms. se in Scandinavia in having a somewhat lar­ The specimens from the Baojing Formation ger average size, tending to have a slightly in western Hunan, China, figured by Zhou et shorter glabella, a pygidium with less cons tric­ al. (1977, PI. 36, figs. 20-21; also Yang 1978, ted posterior axial furrow , slightly larger poste ­ PI.1 , figs. 11-12) as D. incertus differ from ro-Iateral border spines and a less curved Scandinavian specimens primarily in having a posterior margin. He further stated that the more quadratic pygidium with broader borders Scandinavian spec imens tend to lack a delto id and a stouter, posteriorly less acute axis. area. Howeve r, Robison tentatively conc urred Both the figured Chinese cepha lon and pygidi- 62 Frank Nikolaisen & Gunnar Henningsmoen GU · BULL. 419. 1990 um agree very well with those of D. notalibrae (3) The occipital ring has its node (spine) loca­ Opik (1979, pp. 84-86, PI. 19, figs. 1-4, PI. 21, ted very close to the posterior margin. (4)The figs. 1-3) from the Undillan Zone of Goniagnos­ presence of an anteriorly situated palpebro­ tus nathorsti in Queensland, Australia, and ocular ridge. (5) The well-developed epipalpe­ belong most prob ably to that species. bral furrow. (6) The presence of both a postocu­ lar ridge and a metagenal ridge behind the Order Olenellida Resser. 1938 palpebral lobe. The metagenal ridge is seen Family Olenelli dae Vogde s. 1893 to end in a tiny spine in well-preserved speci­ Subfamily Holm iinae Hupe , 1953 mens of Holmia kjerulfi. Kjerulfia lata has at least a slightly protruding angle of the poste ri­ Genus Kjerulfia Kieer, 1917 or margin in the corresponding place. (7) The structure of the hypostome and its broad (tr.) Date of publication. - Kjerulfia and the type and firm connection with the rostraI plate. The species, K. lata, were published by Joh an Kicer hypostome of Holmia kjerulfi has four small, in his paper "The Lower Cambrian Holmia marginal spines post eriorly, as has also (new fauna at Ternten in Norway". The year 1916 observation) Kjerulfia lata. (8) Lack of macro p­ is given on the front page, but on p. 112 it leurae. (9) Lack of obvious opisthothorax.(10) says "Printed April 20th 1917". We consequent­ Presence of longer axial spines on the hinder­ ly regard 1917 as the year of publication, as most axial rings, and thus no single macrospi­ did Poulsen (p. 0193 ) in the part on trilobite ne. systematics in the "Treatise" (Moore, ed., The combination of the above-mentioned 1959), but which is not reflected in its referen­ features is not known in other olenellid gene­ ce list. Some later authors have also given the ra. Thus, Schmidtiellus Moberg, 1906 has lon­ year as 1917. ger (sag.) base of the axial spine of LO, and where the thorax is known , a macrospine and Type species. - Kjerulfia lata Kia:!r, 1917 by an opisthotho rax. Callavia Matth ew, 1897 dif­ original designation. fers i.a. in the shape of the glabella, in the lack of an epipalpebral furrow, and in the Systematic position. -Until Kirer described connection between hypostome and rostral K. lata, material of this form had been referred plate. Neltneria Hupe, 1953 has i.a. a '/ery to Holmia kjerulfi (Linnarsson, 1871) (cf. Kirer different glabella. Wanneria Walcott, 1910 has 1917, p. 71). Kjeru lfia has been assigned to a glabella that resembles those of Holmia and various subfamilies, e.g. to the Holmiinae Kjerulfia, although L3 is not quite as domina­ Hupe, 1953 by Hupe 1953(a) and Repina 1979, ting, but the hypostome is multispinose and to the Callaviinae Poulsen, 1959 by Poulsen the pygidium is bilobed. Bondonella Hupe, 1959 (in the "Treatise") and with doubt to the 1953 differs La. in having an almost parallel­ Neltneriinae Hupe, 1953 of the family Daguinas­ sided glabella, and in not having an -shaped pididae Hupe, 1953 by Bergstrom 1973(c). In L3. the two latter cases, Holmia was retained in the Holmiinae and Holmiidae, respectively. Kjerulfia lata Kieer. 19 17 We are of the same opinion as Ahlberg and Berqstrorn (in Ahlberg et al. 1986, p. 39), who Figs. 3, 7:A-C assigned Kjerulfia to the subfamily Holmiinae (family Holmiidae). Thus, by comparing materi­ 1917 Kjerulfia lata novo gen. & sp. - Kirer, al of Kjeru lfia lata and Holmia kjerulfi, which pp. 73-81, Pis. 9-13, 14, figs. 1-2 (descr. occur associated in Ringsake r, we can subst­ and figs. of all parts of the exoske le­ antiate that Kjerulfia and Holmia are close ly ton). With synonymy list to date. related and should be grouped together in the 1925 Kjerulfia lata Kjer [sic) - Warburg, p. same subfamily. Among the many features in 35, Text-figs. 11, 12a (remarks and copi­ common, we espec ially mention the following: es of Kieer's reconstructions of genicra­ (1) L3 is distinctly wider (tr.) than LO, L1 and nium in dor sal and ventral views). L2 and has its distal ends pointing in a poste­ 1936 Kjerulfia lata Kiaer - Stubblefield, p. re-lateral direction, giving L3 a low (sag., ex­ 422, Fig. 7 (copies of Kieer's reconstructi­ sag.) M-like shape.(2) L1 is less distinctly ons of genicranium in dorsal and ven­ defined distally than the other glabellar lobes. tral views). NGU - BULL. 419, 1990 Lower and Middle Cambrian trilobites 63

1942 Kjerulfia lata Kirer - stermer, pp. Type specimen. - Lectotype (here selected) 137-138, PI. 2, fig. 1 (remarks and fig. is the incomplete and slightly distorted genicra­ of genicranium). nium, PMO 61376, figured by Kirer 1917, PI. 1953b Kjerulfia lata Kiaer - Hupe, Fig. 9A 10, fig. 1, also: Sterrner 1942, PI. 2, fig. 1, (reconstructions of genicranium in dor­ from the Holmia Shale at Ternten Farm, Ringsa­ sal and ventral views). ker, southern Norway. 1959 Kjerulfia lata Kiaer - Harrington, Fig. 28B (dorsal and ventral views of recons­ Finnmark material. - One fragmentary right tructed genicranium). half of a genicranium (PMO 82712). Two very 1959 Kjerulfia lata Kiar, 1917 [sic] - Poulsen, fragmentary external moulds of glabellae (PMO pp. 0193-0194, Fig. 137/1a-b (copies 98549, 98551) may be conspecific. Preserved of Kirer's reconstructions of dorsal exos­ in greenish-grey, micaceous siltstone (with fla­ keleton and ventral view of genicranium). kes of greenish shale) from the middle part 1960 Kjerulfia lata Kiaer - Suvorova (in Tcher­ of the Lower Cambrian massive-bedded quart­ nysheva), Fig. 37 (copy of Kirer's recons­ zite member (D2) of the Doulbasgaissa Forma­ truction of dorsal exoske leton). tion at locality 1. 1965 Holmia sp. - Reading, p. 175 (reported from Finnmark, determination by Hen­ Dimensions. - The genicranium is 33.5 mm ningsmoen). long (genal spine excluded) and approx imately 1970 Holmia sp. - Banks, p. 21 (listed from 66.0 mm wide (twice the width of right half). Finnmark, determination by Hennings­ The two additional glabellae (if conspecific) moen). indicate individuals of approximately the same 1971 Holmia sp. - Henningsmoen, p. 90 (li­ size. sted from Finnmark). 1973c Kjerulfia lata Kirer, - 1916 Bergstrom , pp. 311-312 (remarks). 1974 Holmia sp. - Martinsson, p. 237 (listed from Finnmark). 1978 Holmia sp.- Bjerlykke , p. 49 (listed from Finnmark). 1980 Holmia cf. moberg i - Berqstrorn, p. 12 (mentioned from Finnmark). 1981 Holmia et. mobergi - Bergstrom, p. 22 (listed from Finnmark). 1984b Kjerulfia lata - Ahlberg, p. 19, Fig. 4 (listed). 1984b Holmia cf. mobergi - Ahlberg, pp. 20-21 (Fig. 5), 24 (listed from Finnmark). 1984 Holmia - Henningsmoen, pp. 23-24 Fig. 3. Genicranium of Kjerulfia lata Kiaar, 1917. Reconstruc­ tion based on the incomplete specimens illustrated in Fig. (mentioned from Finnmark and fig. of 7:A-B. PMO 82712 and 98551.Natural size. genicranium). 1985 Kjeru lfia lata Kirer, 1917 - Ahlberg, p. 341 , Fig. 2 (listed). Description (based on PMO 82712).- Genicra­ 1985 Holmia cf. mobergi - Ahlberg, Fig. 3 nium semicircular, twice as wide as long, and (listed from Finnmark). rather low. Cephalic border very narrow (sag.) 1986 Kjerulfia lata - Ahlberg & Berqstrorn (in anteriorly; antero-Iaterally and laterally rather Ahlberg et al.), Fig. 1 (listed). broad and very low; posteriorly narrower and 1986 Holmia et. moberg i Berqstrom , 1973 ­ fairly convex (exsag.), and intersected by a Ahlberg & Bergstrom (in Ahlberg et al.), metagenal ridge which is well delimited by p. 49, Figs. 1, 8 (remarks and figs. of distinct furrows running from SO and from genicrania from Finnmark). posterior end of palpebral lobe. The metage­ 1987 Kjerulfia lata Kirer, 1917 - Nikolaisen, nal ridge may well have ended in a metagenal pp. 305-309, Fig.1 :F-G (remarks and figs. spine (node). Genal spines fairly short , con­ of genicranium). fluent with lateral cephalic border. Genal field posteriorly distinctly wider (tr.) than glabella. 64 Frank Nikolaisen & Gunnar Henningsmoen NGU - BULL. 419. 1990

A very shallow ridge curves outwa rds and gi in having a clearly wider genicranium, wider backwa rds for some distance from the frontal (tr.) genal areas, a less obtuse angle between glabe llar lobe. Glabella laterally defined by frontal glabellar lobe and anterior palpebro­ weak axial furrow, practically missing at L1, ocular ridge, and a less convex (tr.) lateral anter iorly almost conf luent with anterior mar­ borde r. gin of genicranium. L3 wider (tr.) than L1 and In addition to the genotype , Kjerulfia inclu­ L2 and has the shape of a low M. S1 very des only K. selandica Poulsen (1 969, pp. deep and rather broad abaxially, weak adaxial­ 13-16, PI. unnumbered, figs. 1-2) from a bore­ Iy and apparently not transgl abellar; S2 fairly hole near Slagelse, western Zealand, Den­ narrow, transglabellar but not reaching the mark, K. orcina Orl owski (1974, pp. 13-15, PI. periglabellar furrow; S3 narrow, well incised 3, figs. 4-5, PI. 4, figs. 1-8, PI. 5, figs. 1-5; and transglabellar. Palpebral lobes reaching also 1985a, p. 234, text-fig. 2, PI. 1, figs. 5-6) backwa rds to opposite occipital furrow. Anteri­ from the Holmia Zone in G6r Swietokrzyskie or and poste rior palpebro-ocular ridges divided (Holy Cross Mts.), Poland, K. schwarzbachi by a strong epipalpebral furrow (only the proxi­ Ahlberg & Berqst rorn (1986, pp. 51-52, Figs. mal part of the palpebra l lobe is preserved in 10-11) from Lusiatops-Schiefer, Niederlud­ the present material) that continues onto the wigshof, Gorlitz , eastern Germany, and K.? fronta l glabellar lobe to separate L3 and L4 palpebra Ahlberg (1984a, pp. 257-259, Figs. (as defined by Bergst rom 1973b, pp. 209-210, 6-7) from the Holmia kjerulfi-group Zone in the Fig. 1). The epipalpebral furrow is clearly clo­ upper part of the Grammajukku Formation at ser to palpebral area than to extraocular ge­ Delliknas, Leisvall area, central Swedish Lap­ nal area. land. K. lata differs from the Danish form in having much broader (tr.) lateral bord ers, and Remarks. - The present material has been from the Polish and the north ern Swedish referred to as Holmia sp. in most previous form in having a much wider genal field and reports, though none of these reports were lower lateral borders. founded on more than brief studies of it. The An epipalpebral furrow that continues onto Finnmark material, however scarce and frag­ the front al glabellar lobe is distinctly present mentary, agrees in almost all details with the in many olenellid species; from south ern Scan­ description given by Kicer (1917, pp. 73-81) dinavia for instance Wanneria? lundgreni (Mo­ of Kjerulfia lata fro m the Holmia kjerulfi-grou p berg, 1892; see Moberg 1899, PI. 14, fig. 2; zone at Ternten in Ringsaker , southern Nor­ also Berqstrorn 1973c, Fig. 17), H. sulcata way. The most comp lete specimen at hand is Berqstrorn (1973c, pp. 292-294, Figs. 7-8) and slightly smaller than most of Kicer's speci­ H. grandis Kicer (1 917, pp. 70-71, PI. 6, fig. mens. That, together with the difference in 12). However , K. lata seems to differ from preservation, may thus expla in the very small them all in that the epipalpebral furrow is very differences which may be observed , e.g. the close to S3. more rounded frontal glabellar lobe and the lower lateral border. The epipalpebral furrow Order Redlichiida Richter. 1933 cont inues distinctly onto the frontal glabellar Suborder Redlichiina Richter. 1933 lobe, but may have been exaggerated by dor­ Superfamily Ellipsocephalacea Matthew. so-ventral compression. A similar condition is, howeve r, present in the smaller genicranium 1888 figured by Kicer (1917, PI. 9, fig. 2). The Finnmark material has more recently Date ofpublication. - The family name Ellipso­ been referred to as a form possibly conspeci­ cephalidce was introduced by Matthew in his fic with or close to Holmia mobergi Berqst rorn article "Illustrations of the Fauna of the St. (1973c, pp. 288-292, Figs. 3-6) Le. by Berg­ John Group. No. IV". The year 1887 is printed strorn 1980 (p.12), 1981 (p. 22), Ahlberg 1984b on the first page of the article and has since (pp. 20-21, 24) and 1985 (p. 343) and by Ahl­ been attributed to the author's name for the berg & Bergstrom (in Ahlberg et al. 1986, Fig. family and its super- and subfamily derivati­ 8, p. 49). Accord ing to Ahlberg & Bergstrom ves. However, on the front page of the comple­ (198 3, p. 245) H. mobergi belongs to an earli­ te volume, the year of printing for the procee­ er species group of Holmia than does H. kje­ dings and transactions for the year 1887 is rulfi. The Finnmark form differs from H. mober- 1888. For that reason we believe that it is NGU · BULL. 419, 1990 Lower and Middle Cambrian trilobites 65 correct to substitute "Matthew, 1887" with red by Snajdr 1958, PI. 7, fig. 6 (also: Horny "Matthew, 1888". & Bastl 1970, PI. 2, fig. 1).

Family Ellipsocephalidae Matthew. 1888 Finnmark material. - One incomplete cranidi­ Subfamily Ellipsocephalinae Matthew. 1888 um articulated with eleven thoracic segments with counterpiece (PMO 72350, 72351), one Genus Ellipsocephalus Zenker. 1833 very fragmentary dorsal exoske leton with shedded librigenae with counterpiece (PMO Synonym. - ?Strenuella (Ellipsostrenua) Kaut­ 72349, 72348), about twenty-five more or less sky, 1945 (type species by original designati­ fragmentary cranidia of which most are inter­ on: S. (E.) gripi Kautsky, 1945), according to nal moulds or counterparts of such and three Ahlberg & Bergstrom 1978, pp. 12, 17. incomplete thoraces , preserved in mudstone and impure quartzite from the lower Middle Type species. - Elleipsocephalus amb iguus Cambrian quartz ite and shale member (K1) Zenker, 1833 (= subjective junior synonym of of the Kistedal Formation at locality 4. Trilobites Hoffii Schlotheim, 1823) by monoty­ py. Dimensions . - The cranidium articulated with eleven thoracic tergites is 35.5 mm long (the Ellipsocephalus cf. hoffii (Schlotheim. 1823) posteriormost part of thorax and the pygidium is lacking). The cranidium is 15.5 mm long and Fig. 4, Fig. 7:D-1 25.0 mm wide (twice the width of left half). The largest cranidium (internal mould) has an cf . 1958 Ellipsocephalus hoffi (Schlotheim, 18 mm long glabella (excluding the occipital 1823) - Snajdr, pp. 88-92, PI. 7, figs. ring) which suggests a cranidium fully 23 mm 1-8, PI. 8, figs. 1-7, ?8 (descr. and figs. long and 37 mm wide. of topotype dorsal exoskeletons). With synonymy list to date. 1961 An ellipsocepha lid - Henningsmoen, p. 94 (recorded from Finnmark). 1965 Ellipsocephalus sp. - Reading, p. 175 (listed from Finnmark). 1970 Ellipsocephalus hoffi (Schlotheim, 1823) - Horny & Bastl, pp. 166-167, PI. 2, fig. 1 (type specimens listed and fig. of neoty­ pe). 1971 Ellipsocephalid - Henningsmoen, p. 90 (reported from Finnmark). 1974 Ellipsocephalus sp. - Martinsson, p. 237 (reported from Finnmark). 1984 Ellipsocephalus - Henningsmoen, pp. Fig. 4. Cranidium of Eflipsocepha/us ct. nottll (Schlotheim, 23-24 (reported from Finnmark and fig. 1823). Reconstruction based on that of the specimen in of incomplete dorsal exoske leton with Fig. 7:0 . PMO 72350. Approx imately x 2 1/2. shedded librigenae). ct. 1985b Ellipsocepha lus hoffi Schlotheim, 1823 - Orlowski, pp. 252-253, PI. 1, Description . - Cran idium about two-th irds as figs. 1-8 (short descr. and figs. of dor­ long as wide, sub-trapezoidal, but with bluntly sal exoskeleton and cranidia). pointed, mesially somewhat truncated , anterior margin. Preglabellar field short, about as long Type specimen. - Neotype is the practically (sag.) as occipital ring (small specimens have complete dorsal exoskeleton with shedded Ii­ a relatively somewhat longer preglabellar fi­ brigenae preserved in Narodniho Museum in eld). Glabella with subparallel sides, slightly Prague, cat. no. Br-185, from the Zone of widened at base of eye ridges and rounded Hydrocephalus Iyelli (subzone of Ellipsocepha­ in front (but bluntly pointed on internal moulds). Ius hoffii), Jince Formation, at Jince, Cesky Occipital furrow shallow but fairly well pronoun­ (Bohemia), Czechoslovak ia, selected and figu- ced. Occipital ring with a small, low median 66 Frank Nikolaisen & Gunnar Henningsmoen NGU - BULL. 419. 1990

node. Lateral glabellar furrows not visible, Anothe r Bohemian form similar to the pre­ except for a weak ly developed , oblique S1 in sent is the presumably more concurrent E. larger cranidia. Periglabellar furrow well defi­ vetustus Pompeckj (1896, p. 552, PI. 17, fig. ned posteriorly, progressively effaced for­ 3; see Snajdr 1958, pp. 92-94, PI. 6, figs. wards. Fixigenae about as wide as glabella, 21-22, PI. 8, fig. 9, PI. 9, figs. 1-11) from the with faintly developed, narrow and oblique eye Skryje Beds (Zone of Eccaparadoxides pusil­ ridges , and with shallow but distinct posterior Ius), Jince Formation, but which differs in ha­ borde r furrows. Facial sutures rounding off ving cranidia with a broader (sag.) preglabellar antero- Iateral corne rs, subparallet in front of area. eye lobes, slightly convex and gently diverging The cranidia from Finnmark differ from tho­ behind these. Palpebral lobes poorly delimited se of the southern Scandinavian Ellipsocepha­ from palpebral area of fixigenae, about one­ Ius polyto mus Linnarsson (1877, pp. 12-15, quarter as long as cranidium, situated about PI. 2, fig. 1; see westerqaro 1936, pp. 56-58, mid-way between anterior and posterior cor­ PI. 11, figs. 5-7) from the Zone of ners of the cranidium. Surface of cranidium Eccaparadoxides oelandicus in having a wider appea rs smooth. Librigenae, doublure and (tr.) palpebral area of fixigena, a less curved hypostome unknown. Thorax incompletely anterior margin and wider (tr.) pleural areas, known ; one specimen shows twelve tergites. and from the younger Ellipsocephalus lejostra­ Pleurae wider than axis (more so rearwards), cus (Angelin, 1852, p. 24, PI. 19, fig. 3; see and with oblique, moderately broad (exsag.) westeroarc 1950, pp. 11-13, PI. 2, figs. 7-14) pleural furrows. Fulcra slightly further from from the Zone of Ptychagnostus gibbus, which axial furrow than from pleural tips. Anterior has a very similar cranidium, in having non­ pleural band raised into a convex (exsag.) rid­ spinose pleural ends. ge along the anterior margin of pleura. Pleu­ A complete dorsal exoskeleton from Eccapa­ ral ends apparently non-spinose but rather radoxides oelandicus beds in a bore-core from bluntly pointed and bent gently down. Pygidium Bodaharnn, northernmost Gland, Sweden, figu­ poorly known, but apparently small and shor t. red by Wcern (1952, PI. 1, fig. 3) has only twelve thoracic tergites. It differs from the Finn­ Remarks. - The present exoskeletal parts mark form in having much shorter (tr.) pleu­ agree , in as far as the state of preservation rae. Otherw ise its cranidium equals that of allows comparison , rather well with correspon­ Ellipsocephalus polytomus. ding parts in the topo type material (see also A number of both named and unnamed forms Orl owski 1985b, fig. 1) of the presumably from widely scattered areas in central and younger type species, E. hoffii, as described southwestern Europe have been assigned to and figured by Snajdr (1958, pp. 88-92, PI. Ellipsocephalus (e.g. Orlowski 1959b, 1964, 7, figs. 1-8, PI. 8, figs. 1-7, ?8) from the Zone 1965, 1971, 1975a, 1975b, 1985b; Samsono­ of Hydroce phalus Iyelli (subzone of E. hoffii), wicz 1959, 1962; Lendzion 1972; Lendzion in Jince Formation at Konicek, Vystrkov and Jin­ Aren & Lendzion 1978; Sdzuy 1958, 1961, ce in Cesky (Bohemia), as well as with topoty­ 1966, 1968; Linan Guijarro 1978). The state pe mater ial preserved in the Paleontologisk of preservation of most of these , as well as Museum in Oslo. E. hoffii is also recorded and that of the prese nt material, renders compari­ briefly described from both the Zone of Para­ son with the Finnmark form rather difficult and dox ides insuisris. the Zone of P. pinus and speculative, not least due to the fact that ellip­ the Zone of P. po lonicus in the G6r SWi{'tokr­ socep halines have effaced cranidial features zysk ie (Holy Cross Mts .), Poland by Orlo wski and that intraspecific variation is common (1985b, pp. 252-253, PI. 1, figs. 1-8 ). Howe­ (Sdzuy 1961, p. 575). However, some forms ver, the present for m seems to differ from have cranidia quite similar to those present: Bohemian material of E. hoffii in having slight­ E. guerichi Orlowski (1959b, pp. 516-517, PI. ly wider (tr.) fixigenae and correspondingly 1, figs. 6- 10 ; also 1964, p. 82, PI. 4, figs. 4-9; wider (tr.) pleurae, and in having a slightly 1965, p. 137, PI. 1, fig. 6; 1985b, p. 254, PI. more pronounced occipital furrow. It differs 2, figs. 9-15) from the Eccaparadoxides oelan­ from the Polish material, which is preserved dicus Zone of the eastern part of the G6r more like most of the Finnmark material, in SWi ~'tokrzysk ie (Holy Cross Mts .) in centra l having a shorte r (sag.) preg labellar field and Poland differs in having a wider (sag.) pregla­ wider (tr.) fixigenae. bellar area and narrower (tr.) thoracic pleural NGU - BULL. 419. 1990 Lowerand Middle Cambrian trilobites 67

areas; Ellipsocephalus puschiOrlowski (1959b, discussed the matter more than briefly, except p. 517, PI. 2, figs. ta-c: also 1964, p. 82, PI. for Solovyev (1969). He regarded all the gene­ 4, figs. 1-3; 1985b, p. 253, PI. 2, figs. 5-8) from ra above as subgenera and gave amended the same horizon and area differs in having diagnostic concepts. Furthermore, he erected a wider (sag.) preglabellar area; E. leonicus a new subgenus, Eoparadoxides, for the stra­ Sdzuy (1958,p. 238, PI. 1, fig. 5; also 1961, tigraphically low and large species with a flat pp. 575-577 [293-295], PI. 8, fig. 1, PI. 34, fig. and broad (sag., exsag.), sagittally close to 3) from "Trilobite beds", Los Barrios de Luna, anterior border often depressed, anterior crani­ Spain, differs in having a wider (sag.)preglabel­ dial border, transglabellar S1 and S2, narrow lar area and slightly narrower (tr.) fixigenae. (tr.) and small fixigenae, Iibrigenae with a very Finally, it must be noted that the fairly wide broad doublure and a small genal notch, tho­ (tr.) pleurae of the present form are strongly rax of 17-18 tergites, and a small rounded reminiscent of those of Germaropyge germari pygidium. However, it is beyond the scope (Barrande, 1852; see e.g. Horny & Bastl 1970, of this paper to deal closely with the systema­ PI. 2, fig. 2) and G. sanctacrucensis Samsono­ tic classification of Paradoxides s.1. Thus, the wicz (1959, pp. 527, 529, PI. 2, figs. 4-12; see Finnmark material will be treated under gene­ also Orlowski 1975a, pp. 371-372, Pis. 1-4), ric names in accordance with Snajdr (1957; the former from the Bohemian lowermost Midd­ 1958), but accepting the general view that le Cambrian and the latter from the Lower Acadoparadoxides and most probably also Cambrian Protolenus Zone of G6r SWi~okr­ Eoparadoxides are subjective junior synonyms zyskie (Holy Cross Mts.), Poland. Because the of Eccaparadoxides. Iibrigenae of the Finnmark form are unknown, it cannot fully be compared with species of­ Germaropyge. Genus Eccaparadoxides Snajdr, 1957

Superfamily Paradoxidacea Hawle & Corda, Synonyms. - Phanoptes Hawle & Corda, 1847 1847 (nom. trensl. Poulsen 1959, ex Para­ [until ICZN third edition regarded as nom. doxides Hawle & Corda, 1847) obl.] (type species by monotypy: Phanoptes Family Paradoxididae Hawle & Corda, 1847 pulcher Hawle & Corda, 1847 = subjective Subfamily Paradoxidinae Hawle & Corda, synonym of Eccaparadoxides pusillus); Acado­ 1847 paradoxides Snajdr, 1957 (type species by original designation: Paradoxides Sacheri Bar­ Remarks. - The nominate genus was revised rande, 1852); ?Paradoxides (Eoparadoxides) by Snajdr in a preliminary report (1957) and Solovyev, 1969 (type species by original desig­ rather thoroughly soon after (1958). He erec­ nation: Paradoxides Harlani Green, 1834). ted two new genera for species with the ros­ tral plate and hypostome not ankylosed, Ecca­ Type species. - Paradoxides pusillus Barran­ paradoxides and Acadoparadoxides, and awo­ de, 1846 by original designation. ke Barrande's sleeping taxon Hydrocephalus for a group of species with a rather characte­ ristic ontogenetic development. Additional in­ Eccaparadoxides cf. pusillus (Barrande, formation on Paradoxides and Hydrocephalus 1846) was given also later by the same author (1987). These taxa have been somewhat differently Fig.8:F considered by subsequent authors. Some have accepted them as genera (Courtessole 1967; cf. 1958 Eccaparadoxides pusillus (Barrande, Berqstrorn & Levi-Setti 1978), some as sub­ 1846) - Snajdr, pp. 116-128, PI. 20, figs. genera (Sdzuy 1968; Repina 1969; Solovyev 1-46, PI. 21, figs. 1-19, PI. 22, figs. 1-15 1969; Jegorova & Shabanov 1972; Courtesso­ (descr., ontogeny and figs. of ontogene­ le 1973; Lifian Guijarro 1978; Medrano 1982; tic stages and all parts of the holaspid GiI Cid 1984)and some have excluded or rejec­ exoskeleton). With synonymy list to date. ted them (Orlowski 1959a; 1964; 1965; 1985b; non 1962 Paradoxides ct. rugulosus Corda ., Sdzuy 1961; Rushton 1966; Jegorova & Savit­ Hutchinson, pp. 115-116, PI. 23, figs. skij 1969; Jegorova 1976). None of these have 1-3 (remarks and figs. of cranidia). 68 Frank Nikolaisen & Gunnar Henningsmoen NGU·BULL419.1990 cf. 1969 Paradoxides (Eccaparadoxides) pusil­ Remarks. - The present cranidium, however Ius (Barrande, 1846) - Solovyev, PI. 5, very poorly preserved, agrees quite well with figs. 2-3 (reconstructions of meraspid equal-sized cranidia of the presumably stratig­ and holaspid dorsal exoskeleton). raphically concurrent E. pusillus as described cf. 1970 Eccaparadoxides pusillus (Barrande, and illustrated by Snajdr (1958, pp. 116·129, 1846) - Horny & Bastl, pp. 257-258, PI. Pis. 21-22). Nevertheless, it cannot unequivo­ 3, fig. 3 (list of type specimens and fig. cally be included in that species until better of almost complete dorsal exoskeleton and more complete material is available, and fig. by Snajdr 1958, PI. 22, fig. 3) [for is therefore described under open nomenclatu­ synonymous taxa, see Horny & Bastl re. ibid. index p. 343]. The Finnmark form differs from the likewise cf. 1978 Eccaparadoxides pusillus (Barrande) lowermost Middle Cambrian E. insutetts CNes­ - Snajdr, p. 21, PI. 6, fig. 4 (remarks terqard, 1936, pp. 39-40, PI. 7, figs. 19) from and fig. of teratological pygidium). Oland, Sweden (also recorded from central cf. 1978 Paradoxides (Eccaparadoxides) cf. Poland; Orlowskl 1959a; Bednarczyk 1984, pusillus (Barrande 1846) - Llfian Guijar­ and from Rogaland, western Norway; Hen­ ro, pp. 179-180, PI. 9, figs. 1-4 (remarks ningsmoen 1952, PI. 1, figs. 3-4) in having a and figs. of cranidia). less pyriform glabella and a broader (exsag.) cf. 1984 Paradoxides (Eccaparadoxides) pusi­ antero-Iateral cranidial border which is more Ius [sic.] (Barr. 1846) - GiI Cid, pp. pointed sagittally. 309, 311, ?312 (listed, recorded). The present cranidium is also similar to that cf. 1984 Eccaparadoxides pusillus (Barrande, of E. immanis Solovyev (1969, pp. 15-16, PI. 1846) - Snajdr, p. 183 (remarks on recto­ 2, figs. 1-4) from the lower Middle Cambrian type of Phanoptes putcher Hawle & Zone of Oryctocephalops tnschentekii-Schls­ Corda, 1847, PI. 1, fig. 2). tocephalus at Olenek, Anabara in northern cf. 1984 Eccaparadoxides pusillus (Barrande, Jakutsk, northern Siberia, but the latter has a 1846) - Snajdr,p. 189, PI. 14, fig. 2 (re­ less pointed anterior margin and less pronoun­ marks on lectotype of Paradoxides rugu­ ced antero-Iateral borders. losus Hawle & Corda, 1847, p. 32). Two other, geographically widely separated species very similar to E. pusillus are the high­ Type specimen. - Lectotype (selected by ly variable E. eteminicus (Matthew, 1883; see Snajdr 1958) is the complete meraspid dorsal Hutchinson 1962, pp. 114-115, PI. 19, figs. exoskeleton, SBNM coil. Barrande CC 321 3-9, also Martin & Dean 1988, p. 18, PI. 1, No. 1244, figured by Barrande 1872, PI. 9, figs. figs. 1-7, 11-12, 15) from the "Po bennettii 22-23 (also: Snajdr 1958, PI. 20, fig. 43) from Zone" (i.e. the Chamberlain Brooks Formation; the zone with Eccaparadoxides pusillus, Skry­ see Martin & Dean 1988) of southeastern je Beds, Jince Formation, lower Middle Cam­ Newfoundland and E. asturianus (Sdzuy, 1968, brian at the "Podhruskou" locality, near Wfovi­ pp. 91-93, PI. 2, figs. 1-5, ?6, ?13-17) from the ce, Cesky (Bohemia). lower Middle Cambrian of Sebares, eastern Asturias, Spain. E. eteminicus seems to differ Finnmark material. - One incomplete and from E. pusillus in having a somewhat narro­ poorly preserved cranidium (PMO 86583) from wer (sag.) anterior border and a slightly more locality 2, preserved in impure quartzite from pyriform glabella, whereas E. asturianus is the lower Middle Cambrian quartzite and sha­ hardly distinguishable. As it stands, it is quite le member (K1) of the Kistedal Formation. possible that these three forms, together with Three incomplete hypostomes (PMO 86229, some of the other contemporaneous and less 86235, 86540) from locality 4 may belong to well documented species, are conspecific and this form or to Hydrocephalus cf. carens and that the minor differences between them do are described under the latter. not even reflect subspecific features but only intraspecific variation. Dimensions. - The cranidium is 12 mm long and approximately 13 mm wide at eye-line (twi­ Genus Hydrocephalus Barrande, 1846 ce the width of the left half). Synonym. - Phlysacium Hawle & corda, 1847 (type species by monotypy: Phlysacium para- NGU· BULL.419,1990 Lowerand Middle Cambrian trilobites 69 doxum Hawle & Corda, 1847 = subjective juni­ cf. 1984 Hydrocephalus carens Barrande, or synonym of Hydrocephalus carens); Pluto­ 1846 - Snajdr, p. 164 (remarks on lecto­ nia Hicks, 1868 [non Stabile 1864] (type speci­ type of Paradoxides inflatus Hawle & es by monotypy: Plutonia Sedgwickii Hicks, Corda, 1846). 1869); Plutonides Hicks, 1895 (= neonym of cf. 1984 Hydrocephalus carens Barrande, Plutonia Hicks, 1868). 1846 - Snajdr, p. 178, PI. 14, fig. 3 (re­ marks on lectotype of Phlysacium para­ Type species. - Hydrocephalus Carens Bar­ doxum Hawle & Corda, 1847). rande, 1846 by subsequent designation by cf. 1987 Hydrocephalus carens Barrande, Snajdr 1958. 1846 - Snajdr, pp. 101-102.

Hydrocephalus cf. carens Barrande. 1846 Type specimen. - Lectotype is the almost complete internal mould of a meraspid dorsal Fig. 8:A-E exoskeleton, NM L 16579, from the zone with cf. 1958 Hydrocephalus carens Barrande, Eccaparadoxides pusillus, Skryje Beds, Jince 1846, novo emend. - Snajdr, pp. Formation, "Podhruskou" locality, near Wfovi­ 130-139, PI. 24, figs. 1-38, PI. 25, figs. ce, Cesky (Bohemia), figured by Barrande 1-21, PI. 26, figs. 1-11, PI. 27, figs. 1-10, 1852, PI. 49, figs. 9a-b, selected and also figu­ PI. 28, figs. 1 (nonfig. 2 [cf. Snajdr 1987, red by Snajdr 1958, p. 131, PI. 24, fig. 32. p. 101]), PI. 29, fig. 1, PI. 42, figs. 1-3, Finnmark material. - One incomplete cranidi­ 5, 7-8 (descr., ontogeny, figs. of ontoge­ um (PMO 72403), one smaller and very poorly netic stages and all parts of the holas­ preserved cranidium (PMO 86235), three Iibri­ pid exoskeleton). With synonymy list to genae PMO 86227, 86229, 86231), three in­ date. complete but not too poorly preserved hyposto­ 1961 Paradoxides [partim.] - Henningsmoen, mes tentatively assigned to this form (PMO p. 93 (recorded from Finnmark). 86235, 86540, 86229), one fragmentary and 1965 Paradoxides sp. - Reading, p. 175 (li­ very poorly preserved thorax (PMO 86584), sted from Finnmark). one well preserved pleura (PMO 86231) and cf. 1969 Paradoxides (Hydrocephalus) carens some smaller fragments, preserved in shale (Barrande, 1846) - Solovyev, PI. 5, figs. or impure quartzite from the lower Middle 4-5 (reconstr. of meraspid and holaspid Cambrian quartzite and shale member (K1) dorsal exoskeleton). of the Kistedal Formation at locality 4. et. 1970 Hydrocephalus carens Barrande, 1846 - Horny & Bastl, pp. 94-97, PI. Dimensions. - The larger cranidium (PMO 2, figs. 3 (list of type specimens and 72403) is 39 mm long and about 52 mm wide fig. of Hawle & Corda's holotype of at eye-line (estimated by doubling the width Paradoxides dormitzeri) [for synony­ of left half). The largest hypostome (PMO mous taxa, see Horny & Bastl ibid. in­ 86235) is 13.8 mm wide. dex, p. 345]. 1971 Paradoxides sp. - Henningsmoen, p. Description. Cranidium with anterior margin 90 (reported from Finnmark). well rounded. Anterior border narrow and clo­ 1974 Paradoxides sp. [partim.] - Martinsson, se to sagittal line, but slightly widening to­ p. 237 (reported from Finnmark). wards the facial sutures. Glabella pyriform, cf. 1978 Hydrocephalus carens Barrande ­ with occipital ring, L1 and L2 of about equal Snajdr, pp. 10, 15-18, 21, PI. 1, figs. 8-9, length (exsag.). Occipital furrow rather stra­ PI. 4, fig. 1, PI. 6, figs. 5-8, PI. 7, figs. ight. Lateral glabellar furrows shallowest at 7-9, PI. 8, figs. 1-5 (remarks and figs. mesial part; S1 transglabellar and curved slight­ of anomalous parts of the dorsal exoske­ ly inwards and rearwards, S2 gently arched leton). forwards and connected across glabella in a 1984 Eccaparadoxides - Henningsmoen, p. gentle rearwards arch. Internal mould of a 24 (mentioned from Finnmark). smaller cranidium present shows oblique, cf. 1984 Hydrocephalus carens Barrande, strong but short (tr.) S3, whereas the larger 1846 - Snajdr, pp. 151-152 (selection cranidium does not show S3 but faint traces of lectotype of Paradoxides dormitzeri of S4. The post-S2 part of the glabella widens Hawle & Corda, 1847). but slightly forwards and is about three-quar- 70 Frank Nikolaisen &Gunnar Henningsmoen NGU·BULL419,l990

ters as long as the anterior part, which wi­ 6). The difference is, however, trifling and the dens distinctly forwards to mid-length and is three hypostomes are tentatively included he­ well rounded in front. Palpebral area of fixige­ re, because so far only H. et. carens has na about one-third as wide (tr.) as occipital been found at that locality. Further collecting ring, posterior area not preserved in the pre­ at the Digermul peninsula may of course reve­ sent material. Palpebral lobes distinct, gently al that H. cf. carens and E. cf. pusilfus occur curved, diverging rearwards, with base almost together as they do in Bohemia, and that the as far forward as on level with greatest width three hypostomes belong to the latter form. of glabella, and with posterior tips reaching Hydrocephalus is represented in the middle as far back as opposite occipital furrow. Libri­ Middle Cambrian deposits at Andrarum, Sca­ genae (excluding genal spine) about twice as nia in Sweden and at 01eA, Bornholm in Den­ long as palpebral lobe. Lateral and posterior mark. Two forms have been recorded; H. hick­ borders distinct, well defined by sharply inci­ sii palpebrosus (Linnarsson, 1879, pp. 9-11, sed border furrows. Librigenal field nowhere PI. 1, figs. 5-11; see Grenwall 1902; also Berg­ as narrow as border. Genal spine long, cur­ Madsen 1985, p. 370 ) and H. hicksli hicksii ved, at its very point more so slightly in­ (Salter, 1865; see Linnarsson 1883, pp. 14-15, wards. Inner spine angle very slightly obtuse. PI. 3, figs. 1-5; also Grenwall 1902, Berg­ Posterior branch of facial suture shorter than Madsen 1985, p. 360), the latter being the anterior branch; both branches being shorter slightly younger. Both differ from the present than the ocular suture. FHp right-angled. Hypos­ form in having a less rounded anterior rnarqin tome, tentatively assigned to this form, with of the glabella and quite strong S3 and S4. large middle body that tapers quite rapidly The same characters separate the Finnmark backwards, with strong maculae situated far form from the British and Newfoundland speci­ back, and with prosopon made of fine raised es included in Hydrocephalus by Solovyev lines forming a pattern of subparallel inverted (1969, p. 16), Le. H.aurora (Salter, 1869; see U's (cf. below). The quite small thorax present Lake 1935, pp. 212-214, PI. 30, figs. 1-6), H. is effaced and fragmentary, but a well preser­ i1fingi (Lake, 1935, pp. 202-203, PI. 30, fig. 7 ved pleura shows a strongly defined, slightly [? =H. aurora}), H. pervoculus (Howell, 1925, sigmoidal pleural furrow that mid-way is as p. 89, PI. 3, figs. 12-13) and of course H. h. broad (exsag.) as each of the two pleural hicksii (Salter, 1865; see Lake 1934-1935, pp. bands. The pleura extends into a confluent, 196-200, PI. 25, figs. 4-9, PI. 26, fig. 1-2; also curved spine of moderate length pointing obli­ Martin & Dean 1988, p. 19, PI. 3, figs. 4, 7). quely out- and rearwards. Terrace lines dis­ A form worthy to be mentioned here is that tinct on the doublure of the Iibrigenae, elsewhe­ from the northern Siberian Platform described re the surface of the dorsal exoskeleton appe­ and figured by Solovyev (1969, p. 14, PI. 2, ars smooth (the larger cranidium is, however, figs. 5-10; see also Jegorova 1976, pp. 71-72, slightly weathered). PI. 50, figs. 1-2) as Paradoxides (Acadopara­ doxides) sacheri jakutica [sic.], which differs Remarks. - The two incomplete and partly in having a wider palpebral area of fixigenae. poorly preserved cranidia of the Finnmark form The Siberian form may, particularly when the do not allow a very close comparison with larvae or the thorax is found, prove to belong established species, but the Iibrigenae and to Hydrocephalus. well preserved pleura present agree fairly well with equivalent parts of H. carens as illu­ strated by Snajdr (1958, PI. 27, figs. 5, 9, 11-12, PI. 28, figs. 1-2, PI. 29, fig. 1, PI. 30, fig. Genus Paradoxides Brongniart. 1822 1) as well as of topotype material preserved in the Paleontoloqisk Museum in Oslo. On the Synonym. - Vinicella Snajdr, 1957 (type speci­ other hand, the three hypostomes associated es by original designation: Trilobites aestaere­ with the Finnmark material above are apparent­ tus Barrande, 1846 = SUbjective junior syno­ ly somewhat less quadratic in outline than nym of Trilobites gracilis Boeck, 1828; see those of H. carens illustrated by Snajdr (ibid., Snajdr 1978, pp. 24-25). PI. 26, figs. 1-9). Thus, they are slightly more similar to those of E. pusillus (see ibid., PI. Type species. - Entomostracites psrsdoxissl­ 21, figs. 16-18, also; Snajdr 1987, PI. 2, fig. mus Wahlenberg, 1818 (= objective senior NGU-BULL.419,1990 Lowerand Middle Cambrian trilobites 71

synonym of Paradoxides Tessini Brongniart, Wales, selected by Bergstrom & Levi-Setti 1822) by subsequent designation by Barrande (1978) who figured a cast (ibid., PI.2,fig. 1). 1852. Finnmark material. - One cephalon articulated Paradoxides davidis Salter, 1863 (s.l.) with a nearly complete thorax (PMO 72396), one nearly complete young holaspid dorsal Figs. 8:G-I, 9:A-G, 10:A-F exoskeleton (PMO 111.608 with counterpiece PMO 111.609), one thorax (PMO 72358) and 1961 Paradoxides (including P. paradoxissi­ fragments of others, about sixty cranldia, mus) - Henningsmoen, p. 39 (recorded more than twenty larger parts of librigenae, from Finnmark). over twenty hypostomes and five pygidia. All 1965 Paradoxides spp. - Reading, p. 176 (li­ preserved in greenish-grey, shaly mudstone sted from Finnmark). from the Middle Cambrian sandstone and sha­ 1971 Paradoxides paradoxissimus - Hen­ le member (K2) of the Kistedal Formation at ningsmoen, p. 90 (reported from Finn­ localities 5 and 7-13. mark). 1971 Paradoxides cf. davidis - Henningsmo­ Dimensions. - The specimen consisting of a en, p. 90 (reported from Finnmark). cephalon articulated with twenty tergites, is 1974 Paradoxides sp. - Martinsson, p. 237 86 mm long. Fragments of the dorsal exoskele­ (reported from Finnmark). ton indicate individuals of approximately 2.5 1978 Paradoxides - Bj0rlykke, p. 49 (listed times the size of this. The younger holaspid from Finnmark). dorsal exoskeleton is approximately 15 mm 1978 Paradoxides davidis Salter, 1863- Berg­ long (estimated because the anterior of the strorn & Levi-Setti, pp. 6-12, Pis. 2-7, cephalon is missing). The smallest cranidium PI. 8, figs. 1, 3-4, 7-8, PI. 9, figs. 1-3, 5, (PMO 72429) is 4.3 mm long and 6.9 mm wide PI. 10, figs. 1-7 (descr. of phenotypic at eye-line. The largest pygidium present (PMO variation and figs. of holotype and topo­ 72392) is 10 mm long. type material of all parts of the exoskele­ ton). With synonymy list to date. Remarks. - P. davidis has recently been tho­ 1984 to arter av [two species of] Paradoxides roughly revised by Bergstrom & Levi-Setti - Henningsmoen, pp. 23-24 (mentioned (1978)on basis of material from Newfoundland and fig. of early holaspld dorsal exoske­ and from the type locality in South Wales. leton from Finnmark). Including the nominate subspecies, they were 1984 Paradoxides davidis Salter, 1863 ­ able to recognise four different subspecies Whittington, pp. 586-588, PI. 53, figs. founded on variations in the pygidial morpholo­ 1-3,8(remarks and figs. of hypostome). gy and, for one of them, on an abberant num­ 1985 Paradoxides (Paradoxides) davidis davi­ ber of thoracic tergites. Their three new subs­ dis Salter, 1863 - Morris & Fortey, p. pecies, occurring at separate horizons in New­ 110, PI. 7, fig. 3, PI. 8, fig. 4 (figs. of foundland, were thought to have been derived holotype). by allopatric geographic speciation from a stab­ 1988 Paradoxides (Paradoxides) davidis davi­ le stock of the nominate subspecies. A similar­ dis Salter, 1863 - Martin & Dean, p. ly detailed stratigraphical collection is not as 18, PI. 4, figs. 4, 11-17 (remarks and yet available from Finnmark. However, the few figs. of cranidium and pygidia). pygidia at hand indicate that different subspeci­ 1988 Paradoxides davidis Salter, 1863 ­ es occur. The pygidium of the younger holas­ Whittington, pp. 586-588, PI. 53, figs. pid dorsal exoskeleton (Fig. 8:H-I) is rather 1-3, 8, text-fig. 8 (description and figs.of similar to those of P. davidis davidis, another hypostomes). isolated pygidium articulated with the last tho­ racic tergite (Fig. 10:F) strongly recalls that of P. d . trapezopyge, whereas the largest one Type specimen. - Holotype is the external present (Fig. 10:E) does not quite agree with mould of an almost complete but distorted any of those figured by Bergstrom & Levi­ dorsal exoskeleton (BM 45083 and counter­ Setti. The granulation on several fragments, part BM 45084) from Menevian Beds at Porth­ especially such from larger individuals, agrees y-rhaw, St. Davids, Pembrokeshire, South well with that found in P. d. davidis and P. 72 Frank Nikolaisen & Gunnar Henningsmoen NGU-BULL419,1990 d. trapezopyge. The earlier reports of both Order Ptychopariida Swinnerton, 1915 P. paradoxissimus and P. cf. davidis from Finn­ Suborder Ptychopariina Richter, 1933 mark must be viewed in the light of the then Superfamily Ptychopariacea Matthew, 1888 unpublished demonstration of the wide range Family Ptychopariidae Matthew, 1888 of morphological variation in P. davidis as Subfamily Nassoviinae Howell, 1937 well as lack of more complete and better pre­ served material as collected later from Finn­ Genus Nassovia Howell, 1937 mark by Prof. B.-D. Erdtmann and his stu­ dents. Furthermore, the Scandinavian species Type species. - Liostracus globiceps Gron­ of Paradoxides are not yet well enough known, wall, 1902 by original designation. partly because of the scarcity of complete dorsal exoskeletons. It is, for instance, possib­ le that material assigned to P. paradoxissimus Nassovia7 mjol/nir n. sp. may include more than one form. Thus, Strand (1929, p. 349) points out that there are diffe­ Figs. 5a-b, 10:H-J rent morphs of the pygidium. As to the cepha­ Ion, the Finnmark specimens agree very well 1961 an undetermined trilobite - Henningsmo­ with that of P. davidis although the palpebral en, p. 94 (recorded from Finnmark). lobes seem to extend further back than in the 1971 Groenwallia? n.sp. - Henningsmoen, p. material figured by Berqstrorn & Levi-Setti 90 (listed from Finnmark). (1978), which, however, shows some variation 1984 en ukjent trilobitt [an unknown trilobite] (cf. their PI. 5, figs. 3 and 6). - Henningsmoen, p. 24 (mentioned from The cephalic parts of the Finnmark material Finnmark). of P. davidis are also very similar to correspon­ ding parts of the Bohemian P. paradoxissimus Derivation of name. - From the name of the gracilis (Boeck 1828). However, the pattern Old Norse God Thor's hammer, Mj611nir, allu­ of raised lines on the ventral side of the ros­ ding to the likeness of the cranidium to anci­ tri-hypostomal plate agrees with that of P. ent stylised illustrations of the hammer. davidis. Thus, the lateral lines of the hyposto­ me reach the rostraI plate more or less at a Type specimen. - Holotype (PMO 72388) is right angle, whereas they tend to curve more a flattened, slightly longitudinally compressed or less forwards and inwards near the junction but fairly complete cephalon with five articula­ in P. paradoxissimus gracilis. Furthermore, ted thoracic tergites from the Middle Cambrian P. p. gracilis differs in normally having a quite sandstone and shale member (K2) of the Kiste­ narrow (tr.) pygidium where the posterior dal Formation at locality 13. margin is uplifted and slightly pointed, but other morphological types do not seem to be Additional material. - One topotype incomple­ uncommon (cf. Snajdr 1958, PI. 13, figs. 8, te Iibrigena (PMO 72389), and one fragmenta­ 11; 1978a, PI. 5, figs. 2-5, PI. 6, figs. 1-2). ry, most probably conspecific pygidium (PMO Noteworthy is that the tendency to develop a 86143) from locality 16. postero-Iateral spine can be seen in the lar­ gest of the Finnmark pygidia - a spine that Dimensions. - The holotype cephalon is 14 is fully developed in some Bohemian speci­ mm long and 32 mm wide across occipital mens (cf. Snajdr 1958, PI. 11, fig. 1 [Vinicella furrow (restored). desiderata is according to Snajdr 1978, pp. 24-25, synonymous with P. p. gracilis): Horny Diagnosis. - A Nassovia? species with both & Bastl 1970, PI. 3, fig. 4; Snajdr 1978, PI. anterior and posterior branches of facial sutu­ 5, figs. 3-4, PI. 6, fig. 1). Thus, it is possible res strongly diverging, preglabellar area very that P. p. gracilis may occur in Finnmark, but nearly half as broad (sag.) as length of glabel­ further material, especially of complete dorsal la, lateral glabellar furrows weakly impressed, exoskeletons, is needed to verify this. and posterior area of fixigena as wide (tr.) as occipital ring.

Description. - Cephalon subsemicircular, more than twice as wide as long (genal spines NGU - BULL . 419. 1990 Lowerand Middle Cambrian trilobites 73

converging across anterior border to create rounded antero-Iateral cranidial corners (best seen in the facial suture of the associated librl­ gena), posterior branches strong ly diverging slightly sigmoidally backwards (approximately 115 °) and cutting the posterior border rather close to the genal spine. A faint impression on the glabella appears to show the outline of the hypostome. Doublure as wide as bor­ der. Librigena with a medium-sized and very broad-based genal spine. The isolated libri­ gena shows part of the terrace-bearing doublu­ re. The adaxial limitation of it appears to be natural rather than broken, and as it is too b short to reach the axial line of the cephalon, there appears to have been a rostral plate present. The librigena furthermo re shows a very fine radiation of caeacal lines from the eye. The surface of the holotype is less well preserved, but traces of such lines are seen Fig. 5. Nassovia? mjol/nir n.sp. a. Reconstruction of the cephalon with anterior three thoracic tergites based on the on the preglabellar field. Thoracic tergites holotype illustrated in Fig. 10:J. PMO 72388. x 2. b.Recons­ narrow (sag., exsag.), with axial rings apparent­ truction of the pygidium based on the specimen illustrated in Fig. 10:1. PMO 86143. x 3. ly without median nodes, pleurae distinctly wider (tr.) than axial ring, except in first tergi­ te where much shorter, and with fulcra closer to axial furrow than to distal ends. Whether excluded), most probably fairly low, but with pleural ends are pointed or blunt cannot be very deep and broad anterior and lateral bor­ seen in the present material. Pygidium, provi­ der furrows . Cranidium hour-glass-shaped, ded the present specimen is conspecific, trian­ slightly less than half as long as wide and gular in outline and slightly more than twice being widest posteriorly. Preglabellar field very as wide as long. Axis broad (tr.) and compo­ nearly one-third as long (sag.) as glabella. sed of five rings and a terminal axial piece. Anterior border strong , apparently rather con­ Doublure fairly broad. vex (sag.), and only slightly shorter (sag.) than preglabellar field. Glabella gently coniform but Remarks. Nassovia? mjollnir n.sp. seems to markedly truncate (bluntly rounded) anteriorly, occupy an intermediate position between the distinctly defined by narrow but shallow perigla­ monospecific Nasso via and Groenwa l/ia as bellar furrow , and slightly longer than wide. represented by G. microphthalma (Angelin, Lateral glabellar furrows rather shallow and 1852, p. 22, PI. 18, fig. 4; see Westergard only two pairs are discernible in the only crani­ 1953, pp. 32-34, PI. 7, figs. 13-17; also Bruton dium present. Occipital furrow shallow and, 1985, pp. 322-323, Fig. 5:G-I, K-L). The new like lateral glabellar furrows , not reaching the species differs from N. globiceps (Gronwall, periglabellar furrow. Occipital ring one-sixth 1902, pp. 145-146, PI. 4, figs. 12a-b) from the as long (sag.) as glabella, with a faint band Zone of Ptychagnostus punctuosus at 01eA, furrow and apparently with a very low median Bornholm in Denmark, in having the anterior node. Palpebral area of fixigena about half branches of facial sutures strongly divergent as wide (tr.) as width of glabella at eye-line, forwards , glabella less coniform and palpebral posterior area about as wide as occipital ring. lobes closer to the glabella, and from G. mic­ Posterior border furrow (exsag.) but deep and rophthalma from the slightly younger Paradoxi­ almost straight. Posterior border narrow (ex­ des forchhammeri Series of southern Scandi­ sag.). Eye ridges prominent, directed slightly navia in having a shorter preglabellar area, obliquely outwards and backwards . Palpebral more anteriorly situated palpebral lobes, more lobes fairly short (exsag.), situated slightly triangular posterior area of fixigenae, more behind 52. Anterior branches of facial sutures transverse eye ridges and a stronger, much strongly diverging forwards (almost 90°) but less coniform glabella. 74 Frank Nikolaisen & Gunnar Henningsmoe n GU · BULL. 419. 1990

The pygidium differs from that of Andrarina Superfamily Anomocaracea Poulsen . 1927 costata (Angelin, 1854; see Bruton 1985. pp. (nom . tra nsl. Poulse n. 1959. ex Anomocari­ 322-323. Fig. 5:A-D) in having a shorter axis dae Poulsen . 1927) with fewer rings, and from that of G. micropht­ Family Anomocaridae Poulsen . 1927 halma in being more triangular. The gross morph ology of the new species Genus Anomocarina Lermontova. 1940 seems to agree best with that of N. globiceps although close comparison is difficult due to Type species. - Proetus? excavatus Angelin, spars e material of both forms and difference 1852, by original designation. in preservation. For that reason we have inclu­ ded it in Nassovia but with cons iderable Anomocarina7 sp. doubt. The exact stratigraph ical level of the holotype of N.? mjol/nir is uncertain, but beds Fig. 10:K-L yelding a large number of trilobites at other localities on the Digermul peninsula, including Material. - One external mould of a large, the pygidium assigned here, agree stratigraphi­ incomplete pygidium (PMO 72375) and an cally with that of N. globiceps in Bornholm. external mould of a fragment of a slightly lar­ The librigena of N.? mjol/nir has anter ior ger pygidium (PMO 72371), both from the prolongations of the doublure similar to that Middle Cambrian sandstone and shale (K2) found in olenids and in Andrarina (see Bruton of the Kistedal Formation, the former preser­ 1985, p. 323. Fig. 5:J). However. the new ved in micaceous shale from a loose block species differ s markedly from Andrarina in at locality 17, the latter preserved in impure having a much larger preglabellar area, palpe­ mudstone from locality 14. bral lobes situated more posteriorly and clo­ ser to the glabella, and larger, much more Dimensions. - The incomplete pygidium (PMO broad-based genal spines (as in North Ameri­ 72375) has an axial length of approximately can members of Nassov iinae). We agree with 19 mm of which the axis occup ies about 15 Bruton (1985) in that Groenwallia probab ly mm. The width is unmeasurable due to frag­ shou ld be excluded from the Andrarinidae. but mentation, but was possibly about twice the too little is known of the Nassoviinae to inclu­ width. de it there with conf idence. Description. - Pygidial axis taper ing slightly Ptychopariine genus & specie s indet . backwards. composed of six rings carrying strong band furrow s and a large, elevated Fig. 10:G terminal axial piece with a poorly defined, strong ly backwardly tapering projection. Pleu­ Remarks. - A single, 3 mm long, poor ly pre­ ral furrows broad (exsag.). Pleural ribs flat. served and fragmentary external mould of a which gives rise to distinct anterior and posteri­ cranidium occurs assoc iated with Paradoxides or edges . Doublure wide, giving rise to a dis­ davidis, Peronopsis terox sal/esi, Doryagnos­ tinct paradoublural line that is pointed back­ tus incertus and the pygidium here assigned wards sagittally. Middle part of poste rior mar­ to Nasso via? mjol/nir in greenish- grey, shaly gin gently emarginate. mudstone in the sandstone and shale member (K2) of the Kisteda l Format ion at locality 16. Remarks. - The present pygidia are far too It show s a glabella that tapers strongly for­ fragme ntary to allow any close comparison wards and has a very markedly truncate front, with established species. The shape and num­ a moderately broad (sag.) preglabellar field ber of the axis and its rings togethe r with and a quite narrow anterior border. Hence the 'double-edged' pleural ribs may indicate that present cranidium resembles that of UI/aspis they belong to Anomocarina, as exemplified conifron s WestergMd (1948, p. 22, pI. 4, figs. by A. extornata (WestergMd, 1930. p. 17, PI. 14-16) from presumably the Zone of Lejopyge 4, fig. 23; 1950, PI. 3, figs. 12. 15-18, PI. 4, laevigata at Ullavi in Narke, Sweden. but dif­ figs. 3-5) from the Andrarum Limestone at fers in having a broader (sag.) preglabellar Andra rum in Scania, Sweden , A. sibirica (Wes­ field and eyelobes located furth er from the terqard, 1930, PI. 2, figs. 15, 17-18, PI. 3, figs. glabella. 1, ?2) from the upper Middle Cambrian Ben- NGU· BULL.419.1990 Lowerand Middle Cambrian trilobites 75

nett Island, New Siberian islands in the Laptev K2 Sandstone and shale member. 200m. Sea north of Sibiria, and A. splendens Lermon­ Middle Cambrian. tova (1940. PI. 48. figs. 2-2a) from the upper K1 Quartzite and shale member.100m. Midd­ Middle Cambrian of the Anabarsk and Lena­ le Cambrian. Aldansk districts in northern Siberia. Especial­ ly noteworthy is that one of Westergard's figu­ Vestertana Group red pygidia (1950, PI. 4, fig. 4) shows a poin­ Duolbasgaissa Formation (Lower Cambrian): ted paradoubluralline as in the present form. D2Upper Duolbasgaissa member (massive­ bedded quartzite). 300m. However, at least two other species should D1 Lower Duolbasgaissa member (thin­ also be considered as possibly related; Meta­ bedded quartzite). 200-220m. nomocare peta/oides Lermontova (1940, p. 156, PI. 47, figs. 5-5a) from the upper Middle Breivik Formation (Lower Cambrian): Cambrian of Lena, northern Siberia, and B2 Upper Breivik member (green slltsto­ Chondranomocare irbinica Repina (1960, pp. nes), 300-400m. 209-210, PI. 16, figs. 8-9; see Repina et al. B1 Lower Breivik member (green siltstones 1975, p. 150, PI. 22, figs. 13-16) from the lo­ and quartzites. 220-225m. wer Middle Cambrlan of eastern and western Sajan, Kuznetskij Alatau, northern Siberian Stappogiedde Formation: Platform. M. peta/oides differs from the Finn­ S3 Manndraperelv member (red quartzitic mark form in having a well developed postaxi­ sandstone). 180m (With Vendian/Lower Cam­ al ridge and the paradoublural line closer to brian boundary?). the axis, whereas C. irbinica differs in having S2 Innerelv member (blue-green and red­ a longer pygidium with a wider doublure. violet slate). 220-255m. Finally, 'double-edged' pleural ribs are also S1 Lillevatn member (quartzitic sandstone). present in pygidia of Anomocarioides, but 40m. pygidia of that genus differ from that of the Finnmark form in having a long axis with Early Cambrian fossils from the Breivik (B) many more rings. and Duolbasgaissa (D) Formations.

Trilobita: Kjerulfia lata Kicer, 1917 (D2)

Foraminiferida [cf. Glaessner 1963, 1978]: Stratigraphic occurrence of Lower P/atysolenites entiqutsstmus Eichwald, 1860 and Middle Cambrian fossils from (B1) the Digermul peninsula The occurrence in the stratigraphic units is Ichnofossils: given for both trilobites and other fossils. Phycodes pedum Seilacher, 1955 (B1), P. pal­ Some fossils have not yet been described, matum Hall, 1852 (B2), P. sp. (B1), Rusophy­ e.g. hyolithids and several brachiopods. cus (B1, D1, D2), Cruziana (D1, D2), Dimorphi­ cnnus (D1, D2), Diplichnites (D2), Cochlichnus Stratigraphic table. (B1), Plagiomus (D1, D2), Rhizocorallium (D2), Teichichnus (B2), Syringomorpha, determined Mainly from Reading (1965), FC2lyn (1967), here as S. nilssoni (Torell, 1868) (D2), Diplocra­ Banks (1970), Banks et al. (1971) and Welsch terion (D1, D2), Skolithos (D2) and various (1986). other trace fossils. To the above list, mainly from Banks (1970), should be added Bergaue­ Digermul Group (pars) ria (D2) (cf. below), Rusophycus dispar (Lin­ Kistedal Formation (pars): narsson, 1871) (D), recorded by Bergstrom K4 Black shale member. 200m. Upper Cam­ 1981, and the following recorded by Welsch brian. 1983: P/anolites (B1). Peteeopbycus (81) K3 Black quartzite member. 10-35m. Midd­ and Taphrhelminthopsis. Trace fossils have le(?) Cambrian. been recorded as far down as in the middle 76 Frank Nikolaisen & Gunnar Henningsmoen NGU - BULL. 419. 1990 member (S2) of the Stappogiedde Formation The trace fossil Syringomorpha nilssoni (To­ by Reading (1965) and Banks (1970). reil, 1868) is previously known only from Lo­ wer Cambrian beds in southern Sweden (and Remarks. - As pointed out by F0yn & Glaess­ in erratic blocks also in Germany). It occurs ner (1979, p. 27), transitional beds spanning in great numbers in some beds of quartzite the Precambrian-Cambrian boundary should in the upper member (D2) of the Duolbas­ be present on the Digermul peninsula. possib­ gaissa Formation. ly represented by a part of the Manndraper­ A large slab with several large specimens elv member (S3) of the Stappogiedde Formati­ of Bergaueria Prantl, 1946 was collected from on. as discussed by them (I.e.• pp. 29, 45; cf. the same member (D2) by the Oxford Ex­ also F0yn 1985, p. 237). Although an internatio­ pedition to Finnmark 1961 (leader Dr. D.R. nal agreement on the base of the Cambrian Berry). The slab was left at the camp site in has as yet not been reached, the Cambrian Hansvikdalen, but two of the specimens were age of Platysolenites antiquissimus generally brought back. A photograph of the slab and is accepted. The species occurs at several one specimen (PMO 72854) with a diameter localities in northern Scandinavia. Thus, it was around 7cm and vertically sectioned was sent found by S. F0yn in the Breivik Formation to the senior author by Mr. M.F. Tuke. The east of the Digermul peninsula (F0yn 1967; name of the ichnofossil was erroneously given Hamar 1967) and later by N.L. Banks in the as Baueria in the list of fossils by Henningsmo­ same formation on the peninsula itself (Banks en in Reading 1965, Fig. 4 (and in later quota­ 1970, p. 21. Banks 1973), 150m above the top tions of the list). of the Stappogiedde Formation. P. antiquissi­ Early Cambrian and Late Precambrian fos­ mus further occurs in southern Scandinavia sils are known from various localities in nort­ and on the East-European Platform. The speci­ hern Norway and northern Sweden (cf. F0yn es is indicative of the Zone of Platysolenites & Glaessner 1979). Some Early Cambrian and antiquissimus and occurs possibly also in the Late Precambrian acritarchs were reported overlying Zone of Schmidtiel/us mickwitzi as from Finnmark by Vidal 1981. No Early Cam­ discussed by Ahlberg & Berqstrorn (1978), brian (or younger) trilobites have been found Vidal (1981 a, 1981b, p. 192 and 1981c, p. 40) in Finnmark outside the Digermul peninsula. and Ahlberg (1984, p. 4). The occurrence of From the eastern part of the adjoining Troms P. antiquissimus on the Digermul peninsula, county, Vogt (1967) recorded Strenuel/a (Ario­ some 600-700m below the occurrence of Kje­ nel/us) primaeva (Br0gger, 1879) from two ruttie lata, might well indicate the lower part localities. The material was assigned to Stre­ of the Platysolenites range zone as advocated nuaeva inflata Ahlberg & Berqstrorn, 1978 and by F0yn & Glaessner (1979, p. 43) and as to S. inflata? by Ahlberg 1980. From a third suggested by Ahlberg (19 84, p. 5) at least locality in eastern Trorns, Vogt (1967. p. 22) the upper part of the Zone of P. antiquissi­ reported Ellipsocephalus sp. (=Ellipsocepha­ mus. The identification of the olenellid from Ius? sp. Ahlberg 1980). In northernmost Swe­ the middle of the upper member (D2) of the den, Strenuaeva inflata, Comluel/a? lapponica Duolbasgaissa Formation as Kjerulfia lata Ahlberg, 1979, Proampyx triangularis Ahlberg strongly indicates the Zone of Holmia kjerulfi & Bergstrom, 1978 and Ellipsocephalus cf. or the overlying Zone of Proampyx linnarsso­ gripi(Kautsky, 1945) have been described from nl. Thus, Kjerulfia lata is elsewhere known northern Lapland, across the state border from only from these two uppermost. Lower Cam­ eastern Troms. The fauna indicates a late brian zones in Ringsaker, one of the Mj0sa Early Cambrian age (cf. Ahlberg 1985). districts in southern Norway. where its occur­ rence in the Zone of Proampyx linnarssoni Middle Cambrian fossils from K1 ofthe Kiste­ recently has been ascertained (Nikolaisen dal Formation 1987). The presence of K. lata shows that the Duolbasgaissa Formation ranges markedly Trilobita: higher up than in the correlation charts given Ellipsocephalus cf. hoffii (Schlothelm, 1923) by Berqstrorn 1981 (Fig. 1) and Ahlberg 1984 Eccaparadoxides cf. pusillus (Barrande, 1846) (Fig. 1) and which was based on an identificati­ Hydrocephalus cf. carens Barrande. 1846 on of the olenellid as Holmia cf. mobergi Berg­ strorn, 1973. NGU-BULL.419.1990 LowerandMiddleCambrian trilobites 77

Phyllocarida: Doryagnostus incertus (Bmgger, 1878) Undescribed form Paradoxides davidis Salter, 1863 s.l. Nassovia? mjollnir n. sp. Brachiopoda: Ptychopariinae gen. et sp. indet. Dictyonina? sp. (Recorded here, PMO 86240) Anomocarina? sp. Lingulepis? sp. Brachiopoda (Two species described and figu­ Acritarcha (Described by Welsch 1986 from red by Strand 1935): upper part of K1 to 1m above the K1/K2 boun­ Lingulepis sp. Strand 1925: Lingulella (Lingule­ dary): pis) cf. roberti Matthew, 1895 Aranidium granulatum Welsch, 1985 Other inarticulates Cristallinium cemtxtense (Slavikova, 1968) Billingsella sp. Strand 1935: B. cf. retrottexe C. ovlttense (Cramer & Diez, 1972) (Matthew, 1896). Figured also by Erdtmann Eliasum cf. E. lIaniscum Fombella, 1977 1983, p. 20. Micrhystridium M. breviacanthum Slavikova, 1968? Hyolithida (Material not yet descibed. One M. brevicornum Jankaukas, 1976 specimen figured by Erdtmann 1983, p. 21). M. dissimilare Volkova, 1969 M. lanatum Volkova, 1969 Varla: M. tuoomtense Kirjanov, 1974 Hyolithellus? sp. M. obscurum Volkova, 1969 Problematicum M. semiapertum Welsch, 1986 M. shinetonense Downie, 1958 Acritarcha (Described by Welsch 1986 from M. spinosum Volkova, 1969 middle part of K2): M. stellatum Deflandre, 1945 Cristallium cambriense (Slavikova, 1968) M. tornatum Volkova, 1968 C. ovil/ense (Cramer & Diez, 1972) Multiplicisp nseridtum martae Cramer & Diez, Eliasum cf. E. lIaniscum Fombella, 1977 1972 Micrhystridium aft. M. breviacanthum Slaviko­ va, 1968? Ichnofossils (Horizontal and vertical burrows M. obscururn Volkova, 1969 recorded by Reading 1965) M. stellatum Def/andre, 1945 Timofeevia lancarae (Cramer & Diez, 1972) Remarks. - The trilobites occur near the base T. cf. T. microretis Martin, 1981 of K1 (Reading 1965, p. 181). The trilobite T. pentagonalis (Vanguestaine, 1974)? fauna in K1 resembles lower Middle Cambrian T. phosphoritica Vanguestaine, 1978 faunas in Czechoslovakia and Poland rather than in Scandinavia. The two paradoxidid spe­ Ichnofossils (Horizontal burrows recorded by cies suggest the lowermost Middle Cambrian Reading 1965.) zone in the Bartandian area in Czechoslova­ kia, Le. the Zone of Eccaparadoxides pusil/us, Remarks. - As far as there is stratigraphical probably corresponding to the Zone of Eccapa­ control, the trilobites collected from K2 come radoxides lnsulsrts of the zonal group of Ecca­ from the lower half of the member. The upper paradoxides oelandicus in Scandinavia. Ellip­ part of K2 has yielded non-trilobite fossils, socephalus hoffii occurs in Bohemia in the predominantly brachiopods. According to middle of the Middle Cambrian, but is recor­ Strand (1935), the two brachiopod species ded in Poland also from the lower Middle that he described suggest the zonal group Cambrian Zones of E. insularis and E. pinus of Paradoxides forchhammeri. However, the (et. Orlowski 1985b). Middle Cambrian brachiopod faunas from the Kistedal Formation are in need of a revision Middle Cambrian fossils from K2 ofthe Kiste­ now that so much more material has been dal Formation collected. The trilobite fauna known from K? may safe­ Trilobita: ly be assigned to the Zone of Ptychagnostus Peronopsis terox (Tullberg, 1880) sallesi (Ber­ punctuosus in the zonal group of Paradoxides geron, 1889) paradoxissimus. Thus, Doryagnostus incertus 78 Frank Nikolaisen & Gunnar Henningsmoen NGU·BULL419.1990

and Paradoxides aeviais are recorded elsew­ Lower Cambrian here in Scandinavia only from the Zone of Zone of Holmia kjerulfi or, possibly, the overly­ P. punctuosus. The symbol of this zone is ing Zone of Proampyx Iinnarssoni (in the midd­ 1co, in Norway and B4 in Sweden. Doryagnos­ le of 02). tus incertus is known with certainty only from The number of zones is small as compared Scandinavia. Paradoxides davidis occurs both to the standard succession of southern Scandi­ in Scandinavia, Britain, Newfoundland, Nova navia. This may not necessarily be due to Scotia and possibly in Spain. In Britain and gaps, since the fossils are mainly concentrated Newfoundland it occurs in horizons correspon­ in the argillaceous horizons in a predominant­ ding to B4 and the underlying Zone of Hypag­ ly arenaceous succession. Furthermore, in nostus parvifrons (1 C"f2' B3) (Bergstrom & Levi­ some shales no fossils have been found, in Setti 1978, p. 15), but for Britain restricted to others only non-trilobite fossils. The lack of the zone of P. punctuosus only by Thomas et calcareous concretions or beds on the Diger­ al. (1984, p. 11). Peronopsis ferox sal/esi, not mul peninsula is in contrast to the alum shale earlier recorded from Scandinavia, occurs in sequence in southern Scandinavia, where fos­ middle Middle Cambrian horizons in France sils generally are well preserved in the limesto­ and Spain. Berg-Madsen (1985, 1986) has gi­ nes, but in many localities are missing or rare ven good reasons for including the Scandinavi­ in the shale (Andersson, Oahlman, Gee & an Zone of Ptychagnostus lundgreni - P. Snail 1985, p. 35).

nathorsti (1coH C1) in the Zone of P. punctuo­ sus (1co" B4). Whereas the former zone was earlier referred to the zonal group of Paradoxi­ Affinites of the trilobite faunas des forchhammeri in Sweden (cf. e.g. Wester­ Lower Cambrian gard 1946, p. 8), it had already been assigned The Early Cambrian trilobite found, Kjerulfia to the underlying zonal group of P. paradoxissi­ lata, is elsewhere known only from southern mus in Norway (Henningsmoen 1956). Berg­ Norway. It may be added that the characteris­ Madsen's concept of the Zone of P. punctuo­ tic trace fossil Syringomorpha nitssoni is else­ sus (now 1co, B4-C1) in Scandinavia allows where known in situ only in southern Sweden. correlation with the North American P. punctuo­ sus Interval-zone, established by Robison Lower Middle Cambrian (1984, p. 5). The trilobites in the lower member (K1) of the Kistedal Formation belong to genera which Middle(?) Cambrian Member of the Kiste­ occur elsewhere in Scandinavia, but the speci­ dal Formation K3 es are different and are all compared to speci­ es in Czechoslovakia and Poland (p. 77). Only trace fossils (Cruziana) have been found in the black quartzite member (K3) of the Kiste­ Middle Middle Cambrian dal Formation. The size of the specimens of The trilobite species in the second member Cruziana (almost 3 cm wide) suggests a Midd­ (K2) of the Kistedal Formation are likewise le Cambrian age, since the known Upper Cam­ assigned to genera known elsewhere in Scan­ brian trilobites are small in this area. dinavia. Doryagnostus incertus is known with certainty only from Scandinavia, whereas Para­ doxides davidis occurs in Scandinavia, Britain, Newfoundland, Nova Scotia and possibly in Spain. One species, Nassovia? mjollnir;s new, Trilobite zones and the subspecies Peronopsis ferox sal/esi The trilobite faunas found in the Lower and has only been recorded from Spain and France. Middle Cambrian beds on the Oigermul penin­ The Middle Cambrian trilobite faunas in K1 sula indicate the following zones. and K2 appear more meagre than in contempo­ raneous faunas in southern Scandinavia. This Middle Cambrian may partly be due to the long research history Zone of Ptychagnostus punctuosus (within the in the numerous outcrops in southern Scandi­ lower part of K2) navia and also to the lack of calcareous sedi­ Zone of Eccaparadoxides pusillus/ E. insularis ments in K1 and K2. However, this does not (near the base of K1). explain the occurrence in the K1- and K2- NGU- BULL.419,1990 Lower and Middle Cambriantrilobites 79 faunas of trilobites not known from the Alum Acknowledgements Grateful thanks are due to Dr. Harold G. Reading and his Shale Formation in southern Scandinavia. A collaborators (Oept. of Geology and Mineralogy, Oxford corresponding situation continues in the Upper University) for presenting fossils collected by them and Cambrian and Lower Tremadoc trilobite (ole­ manuscript data to the Paleontologisk museum, and to Dr. Bernd-Oietrich Erdtmann (then at Geologisch-Paleontologi­ nid) faunas of the Oigermul peninsula, as no­ scnes Institut, University of Gottingen) for presenting the ted by Nikolaisen & Henningsmoen (1985), trilobites collected by him and his coworkers. The writers express grateful thanks to Nansenfondet for who suggested that the differences in faunas grants for their two collecting trips to the Oigermulpeninsu­ might be due to the severe bottom conditions la, and to Amanuensis Kari E. Henningsmoen (Institute of Geology, University of Oslo) for assistance in the field in prevailing in the 'Alum Shale' sea in southern 1963. Scandinavia. Great thanks are due to numerous persons in the Tana area for hospitality and assistance with boat transports. Two anonymous referees are also thanked for several suggestions which improved the manuscript. Hans-Arne Nakrem is thanked for help with PC techniques. 80 Frank Nikolaisen & Gunnar Henningsmoen NGU-BULL.419.1990

Barrande, J. 1852: Systeme Silurien du centre de la BoM­ References me. lere Pertie. Recherches paleontologiques, 1. Texte - crusteces: Trilobites. xxx + 935 pp, 51 Pis. Prague­ Ahlberg, P. 1979: Early Cambrian trilobites from Mount Paris. (Not seen.) Luopakte, northern Sweden. Sver. geol. Unders. C Barrande, J. 1872 : SysMme Silurien du centre de la BoM­ 765, 12 pp. me. tore partie. Recherches pal6ontologiques, Supple­ Ahlberg, 1980: Early Cambrian trilobites from northern Scan­ ment au Vol 1. Trilobites, crustsces et Poissons. xxx dinavia. Nor. Geol. Tidsskr. 60, 153-159. + 647 pp, 35 Pis. Prague-Paris. (Not seen.) Ahlberg, P. 1984a: Lower Cambrian trilobites from the Lais­ Basselt, G. & Cocks, L.R.M. 1974: A review of Silurian vall area, northern Sweden. Geol. Poren. Stockh. For­ brachlopods from Gotland. Fossils & Strata 3, 56 pp, handl. 105 [for 1983], 251-259. 7 Pis. Ahlberg, P. 1984b: Lower Cambrian trilobites and biostratig­ Bednarczyk, W. 1984: Biostratigraphy of the Cambrian raphy of Scandinavia. Lund Publ. Geol. 22, 38 pp. Lund. deposits in the Leba area. Acta Geol. Polonica 34, Ahlberg, P. 1984c: A Lower Cambrian trilobite fauna from 95-110, 6 Pis. Jarnttand, central Scandinavian Caledonides. Geot. Fo­ Bergeron, J. 1888: Note sur la presence de la Faune primor­ ren. Stockh. Forhandl. 105 [for 1983], 349-361. ctate (Paradoxidien) dans les environs de Ferrals-Ies­ Ahlberg, P. 1985: The Lower Cambrian trilobite faunas from Montagnes (Herault). Bull. Soc. Geot. France, Ser. 3, the Scandinavian Caledonides - a review. In Gee. D.G. 16 [1887-1888],282-285. & Sturt, BA (eds.) The Caledonide Orogen Scandina­ Bergeron, M.J. 1889: Etude geologique du Massif Ancien via and Related Areas. John Wiley & Sons Ltd., Chiches­ situe au sud du plateau central. Ann. Sci. Geol. 22, iv ter, 339-346. + 361 pp, 9 Pis. Ahlberg, P. & Bergstrom, J. 1983: Lower Cambrian trilobi­ Berg-Madsen, V. 1985: The Middle Cambrian of Bornholm, tes from southern Swedish Lapland. Geol. Foren. Denmark: A stratigraphical revision of the lower alum Stockh. Forhandl. 104 [for 1982], 241-246. shale and associated anthraconites. Geol. Foren. Ahlberg, P., Bergstrom, J. & Jchansscn, J. 1986: lower Stockh. Forhand1. 106 [lor 1984). 357-376. Cambrian olenellid trilobites from the Baltic Faunal Pro­ Berg-Madsen, V. 1986: Oversigt over det reviderede Mellem vince. Geol. Foren. Stockh. Forhandl. 108, 39-56. Kambrium pA Bornholm. Oansk geol. Foren., Arskr. f. Anderson, A., Dahlman, B., Gee, D.G. & Snail, S. 1985: 1985, 1-13. The Scandinavian Alum Shales. Sver. geol. Unders. Bergstrom, J. 1970: Rusophycus as an indication of early Ca 56, 50 pp. Cambrian age. In Crimes, T.P. & Harper, J.C.(eds.). Angelin, N.P. 1852: Pa/reont%gica Svecica. Pars I; Iconog­ Trace fossils. eeot. J., Spec. 155. 3, 35-42. raphia crustaceorum tormettonis transitionis. Fasc. " Bergstrom, J. 1973a: Organization, life and systematics of 1-24, Pis. 1-24. Holrnlze (Stockholm). (Not seen, quoted trilobites. Fossils and Strata 2, 69 pp, 5 Pis. Oslo. Angelin 1878.) Bergstrom, J. 1973b: Trilobite segmentation. Acta Geol. Angelin, N.P. 1854: Pa/reont%gia Scandinavica. Pars I; Polonica 23, 207·219. Crustacea formationis trsnsitionis. Fasc. 2, i-ix, 21-92, Bergstrom, J. 1973c: Classification of olenellid trilobites and Addenda & Corrigenda 1p, Pis. 25-41 [new pis. 20 and some Balto-Scandian species. Nor. Geol. Tidsskr. 53, 22 for the above paper]. Holrnlee (Stockholm). (Not 285-314. seen, quoted Angelin 1878.) Bergstrom, J. 1980: The Caledonian margin of the Fenno­ Angelin, N.P. (ed. G. Lindstrom) 1878: Pa/reontologia Scandi­ scandian shield during the Cambrian. In Wones, D.R. navica. Pars. I; Crustacea formationis transitionis. Fasci­ (ed.). The Caledonides in the U.S.A. (Proc. IGCP pro­ culi I & 11. Pp. i-ix, addenda et corrigenda 1 p, 1·24, ject 27: Caledonide Orogen, 1979 meeting, Blacksburg. 21-91, Pis. 1-41, appendix 93-96 and pis. " 1a, 2-3 Virginia) Oept. Geol. Sci., Virginia Potytecnn. Inst. and and 42 from Angelin's MS. Holrnlee (Stockholm). State Univ., Mem. 2, 9-13. Areri, B. & Lendzion, K. 1978: Charakteristyka stratygraficz­ Bergstrom, J. 1980: Middle and Upper Cambrian biostratig­ no-litologiczna wendu i kambru dolnego (Stratigraphic­ raphy and sedimentation in south central Jlimtland, Iithological cnaractertsttcs of the Vendian and Lower Sweden. Geol. Foren. Stockh. Forhandl. 102,373-376. Cambrian). Prace Inst. Geol. 90, 7·49, 3 Pis. Praha. [In Bergstrom, J. 1981: Lower camortan shelly faunas and Polish with English summary.] biostratigraphy in Scandinavia. In Taylor, M.E. (ed.). Banks, N.L. 1970: Trace fossils from the late Precambrian Short Papers for the Second International Symposium and Lower Cambrian of Finnmark, Norway. In Crimes, on the Cambrian System 1981. U.S. Geol. Surv. Open­ T.P. & Harper, J.C. (ads.). Trace Fossils. Oept. Geol. File Rept. 81-743, 22-25. Miner. Univ. Oxford, N.S. Publ. 426,. 19-34. Bergstrom, J. & Ahlberg, P. 1981: uppermost Lower Cam­ Banks, N.L. 1973a: Trace Fossils in the Halkkvarre Section brian biostratigraphy in scanta, Sweden. Geol. FOren. of the Dividal Group (7late Precambrian - Lower Cam­ Stockh. Forhandl. 103, 193-214. brian), Finnmark. Nor. geol. unders. 288, 1-6. Bergstrom, J. & Levi-Selti, R. 1978: Phenotypic variation Banks, N.L. 1973b: Innerelv Member: Late Precambrian in the Middle Cambrian trilobite Paradoxides davidis Marine Shelf Deposit, East Finnmark. Nor. geol. un­ Salter at Manuets, SE Newfoundland. Geol. Palaeon­ ders. 288, 7-25. tOI. 12, 1-40 Banks, N.L., Edwards, M.B., Geddes, W.P., Hobday, D.K. Beyrich, E. 1845: Ueber einige bOhmische Trilobiten. (Hi) + & Reading,H.G. 1971: Sedimentation, regional stratigrap­ 47 pp, 1 PI. G.Reimer, Berlin. hy and correlation. Late Precambrian and Cambro­ Bj0rlykke, K. 1978: The eastern marginal zone of the Caledo­ Ordovician sedimentation in East Finnmark. Nor. geol. nide Orogen in Norway. In Schenk, P. (ed.). Caledonian­ unoers. 269, 197-236, 6 Pis. Appalachian Orogen of the North Atlantic Region. Ge­ Barrande, J. 1846. Notice preliminaire sur le Systeme Siluri­ 01. Surv. Canada, Paper 78-13, 49-55. en et les Trilobites de BoMme. vi + 97 pp. Leipsic. Boeek, C. 1828'. Notitser til Lreren om Trilobiterne. Mag. (Not seen.) f. Naturvid. 8[for 1827], 11-44, 1 PI. Christiania (Oslo). NGU- BULL.419,1990 Lowerand Middle Cambrian trilobites 81

Bmgger, W.C. lB7B: Om paradoxidesskifrene ved Krekling. hie der bOhmischen Trilobiten. 176 pp, 1 chart, 7 Pis. Nyt Mag. f. Naturvid. 24, 1B-BB, 6 Pis. Kristiania (Oslo). J.G. caive'scne Buchhandlung, Prag, also 1848, Ab­ Brongniart, A. 1B22: Des corps organises fossiles nomrnes handl. kgl. bonm. Gesell. Wisensch., Ser. 5, 5, 117-292, Trilobites. In Brongniart, A. & Desmarest, A.-G. Histoire 1 chart, 7 Pis. Prag. naturelle des crusteces tossnes, sous les rapports Henningsmoen, G. 1951: Remarks on the classification of zoologiques et geo1ogifW8S. F, G. Levrault, Paris, 1-65, trilobites. Nor. Geol. Tidsskr. 29, 174-217. Pis. 1-4. Henningsmoen, G. 1952: Early Middle Cambrian fauna from Bruton, D.L. 19B5: Trilobita. In Bruton, D.L., Harper, D. Rogaland, SW Norway. ( Paradoxides oelandicus stage A.T., Gunby, l. & Naterstad, J. Cambrian and Ordovi­ c lea). Nor. 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Assoc. 1868, 68-69. 5 Pis. Hicks, H. 1869: Rept. Brit. Assoc. 1869. (Not seen.) F0yn, S. 1967: Dividal-gruppen l"Hyolithus-sonenj I Finn­ Hicks, H. 1B71: Descriptions of New Species of Fossils mark og dens tornotd til de eokambnske-kambrtske from the Longmynd Rocks of St. David's. In Harkness, formasjoner. Nor. geol, unders. 249, 1-84, 2 PIs. R & Hicks, H. On the Ancient Rocks of the St. Da­ Feyn, S. 19B5: The Late Precambrian In northern Scandina­ yid's Promontory, South Wales, and their Fossil Con­ via. In Gee D.G. & Sturt, BA (eds.). The Caledonide tents. Q. JI geol. Soc. Lond. 27, 399-404, Pis. 15-16. Orogen - Scandinavia and Related Areas. John Wiley Hicks, H. 1872: On some Undescribed Fossils from the & Sons Ltd., Chichester, 233-245. Menevian Group. Q. J. geol. Soc. Lond. 28, 173-185, F0yn, S. & Glaessner, M.F. 1979: Platysolenites, other ani­ Pis. 5-7. mal fossils, and the Precambrian-Cambrian transition Hicks, H. 1895: On the Genus Plutonides (non Plutonia) in Norway. Nor. eeo; Tidsskr. 59, 25-46. from the Cambrian Rocks of St. David's. Geol. Mag. Fritz, W.H. 1973: Medial Lower Cambrian trilobites from the (N.5.), Dec. 4, 2, 230-231. Mackenzie Mountains, northwestern Canada. Geol. Holm, G. 1890: Forsteningar frAn Lappland, insamlade af Surv. Canada, Pap. 73-24, 43 pp, 1 chart. E. Mortsell. Geol. Foren. Stockh. Forhandl. 12,259-267, GiI Cid, D. 19B1: Los Trilobites Agn6stidos del camonco also Sver. geol. Unders. C 115, 5-13. Inferior y Medio de Espana. Bol. Geol, Miner. 92-2, Horny, R & Bastl, F. 1970: Type Specimens of Fossils in 111-126. the National Museum, Prague. Volume 1. Trilobita. 354 GiI Cid, D. 19B4: El genero Paradoxides en Espafia ; sus pp, 20 Pis, 4 tables. Museum of Natural History, Prague. especies y posici6n estragrafica (Parte I). Z. dt. geol. Howell, B.F. 1925: The Faunas of the Cambrian Paradoxi­ Ges. 135, 307-315. des Beds at Manuels, Newfoundland. Bull. Amer. 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(Not Ortowski, S. 1965: Rewizja fauny kambru llrokowego z seen.) g6ry Slowiec (G6ry $wi~tokrzyskie) (A revision of the Matthew, G.F. 1888: Illustrations of the Fauna of the SI. Middle Cambrian fauna from the Slowiec hill, Holy Cross John Group, No. IV. Part I. Description of a New Speci­ Mts.). Biul. Geol. Univ. Warszawa 6, 3-165, 7 Pis. [In es of Paradoxides (Paradoxides regina). Part 11. The Polish with extensive English summary pp. 134-146.] Smaller Trilobites with Eyes (PtychoparidaJ and Ellipso­ Ortowski, S. 1971: The Middle Cambrian of the Klimont6w cephalidaJ). Proc. Trans. R. Soc. Canada 5, sect. 4 [for anticlinorium, Holy Cross Mts. Acta Geol. Polon. 21, 1887], 115-166, PIS. 1-3. 349-359, 1 PI. Matthew, G.F. 1897: What is the Olenellus Fauna? Amer. Ortowski, S. 1974: Lower camortan biostratigraphy in the Geol. 19, 396-407. (Not seen, quoted Walcott 1910.) Holy Cross Mts, based on the trilobite family Olenelli­ McNamara, K.J. 1978: Paedeomorphosis in Scottish olenel­ dae. Acta geol. Polon. 24, 1-16, Pis. 1-6. lid trilobites (Early Cambrian). Palaeontolgy21,635-655. Ortowski, S. 1975a: The systematic position and ontogeny Martin F. & Dean, w.r. 1988: Middle and Upper Cambrian of the Lower Cambrian trilobite species Ellipsocephalus acritarch and trilobite zonation at Manuels River and sanctacrucensis (Samsonowicz, 1959). Acta geol. Po­ Random Island, eastern Newfoundland. Geol. Surv. Ion. 25, 369-375, 4 Pis. Canada Bull. 381, i-vi, 1-91. Orlowski, S. 1975b: Lower Cambrian trilobites from Upper Medrano, T,P. 1982: El cambrico entre Viniegra de Abajo Silesia (Goczalkowice borehole). Acta geol. Polon. 25, y Mansilla (SIerra de la Demanda, Logrono). Trilobites 377-383, 2 Pis. e Ichnofosiles. Biblioteca de Temas Riojanos, Instituto Ortowski, S. 1985a: Lower Cambrian and its trilobites in de estudios RiojanOS, Logrono 1982. the Holy Cross Mts. Acta geol. Polon. 35, 231-250, 7 Pis. Moberg, J.Chr. 1892: orn Olenellusledet i sydliga Skandina­ Ortowski, S. 1985b: New data on the Middle Cambrian trilo­ vien. Forhandl. ved de skandin. naturforskeres 14. bites and stratigraphy in the Holy Cross Mts. Acta meae; 434-439. Kj"benhavn (Copenhagen). geol. Polon. 35, 251·263, 7 Pis. Moberg, J.Chr. 1899: Sveriges alsta kanda trilobiter. Geol. Pompeckj, J.F. 1896: Die Fauna des Cambrium von Tej­ Foren. Stockh. Forhandl. 21, 309-348, Pis. 13-15. rovic und Skrej in Bohmen. Jb. d. k. k. geol. Rei­ Moberg, J.Chr. 1906: Footnote, p. 35 in Moberg, J.Chr. & chanst. 1895, 45, 495-614, Pis. 13-17. Segerberg, C.O. Bidrag till kannedornen om Ceratopyge­ Poulsen, Chr. 1927: The Oarnbrian, Ozarkian and Canadian regionen med sarskild hansyn till dess utveckling i Faunas of Northwest Greenland. ME'ddel. Gr"nland 70, FogelsAngstrakten. Meddel. Lunds geol. f/iltkl., Ser. B, 233-343, Pis. 14-21. (Not seen.) 2, also Kongl. Fyslogr. S/illsk. Handl. [N.F.] 17, 116 pp, Poulsen, Chr. 1959: Olenellidae. In Moore, RC. (ed.). Treati­ 7 Pis. se on Invertebrate Pa/eontology. Part O. Arthropoda I. Moore, RC. (ed.) 1959; Treatise on Invertebrate Paleontolo­ Geol. Soc. America and univ. Kansas Press. Lawren­ gy. Part O. Arthropoda I. xix + 560 pp. Geol. Soc. ce, Kansas, 0191-0198. America and Univ. Kansas Press. Lawrence, Kansas. Poulsen, Chr. 1969: The Lower Cambrian from Slagelse Morris, S.F. & Fortey. RA. 1985: Catalogue of the type no. 1, Western Sealand. Danm. Geol. Under., 2. Reek­ and figured specimens of Trilobita in the British Muse­ ke, 93, 27 pp, 1 PI. um (Natural History). 183 pp, London. Reading, H.G. 1965: Eocambrian and Lower Palaeozoic Munier-Chalmas, E.P. & Bergeron, J. 1888: Etude paleonto­ geology of the Digermul Peninsula, Tanafjord, Finn­ logique. In Bergeron, J. Sur la presence de la faune mark. Nor. geol. unders. 234, 167·191. prtmordlale (ParadOxidien)dans les environs de Ferrals­ [Repina, L. N.] Penaaa, JL a 1960: RoMIlJIeKCbI les-Montagnes (Herault). Compt. Rend. I'Acad. Sci. TPIlJIOOIlTOB HIlMHerO 11 CpeJIHero KeMOpllH 106, 375-377. 3aITSJIHOll qaCTIl BOCTOqfiOrO C8l'IHa Nikolaisen, F, 1987: Olenellid trilobites from the uppermost [The composition of trilobites in the Lower and Middle Lower Cambrian Evjevik Limestone at Temten in Ringsa­ Cambrian in the western part of East Sajan]. Regio­ ker, Norway. Nor. Geol. Tidsskr. 66[for 1986], 305-309. nal. Stratigr. SSSR 4, 171-236, 19 Pis. Akad. Nauk Nikolaisen, F. & Henningsmoen, G. 1985: Upper Cambrian SSSR, Moscow. [In Russian.] and lower TremadOC olenid trilobites from the Digermul [Repina, L. N.] PeullHa, JL a 1969: peninsula, Finnmark, northern Norway. Nor. geol. un­ ders. Bull. 400, 1-49. TpllJIoOllTbI HllMHerO 11 CpeJIHero Opik, AA 1967: The Mindyallan Fauna of North-Western KeMOPllll ora CIlOIlPIl (Ha cetorellcTBo Queens land. Bull. Bur. Min. Resour., Geol. Geophys., Redlichloidea). llacTi> 11 Austr. 74, 1 (text), xvi + 404 pp, 2 (appendices, plates, [Lower and Middle Cambrian trilobites of southern Sibe­ index), v + 167 pp. ria (Superfamily Redlichioidea). Part 2]. Akad. Nauk Opik, AA 1979: Middle Cambrian agnostids: Systematics SSSR, Sibir. otdel., Trud. Inst. geol. geofiz. 67, 112 pp, and biostratigraphy. Bull. Bur. Min. Resour., Geol. Geo­ 4 Pis. Izdat. "Nauka", Moscow. [In Russian.] phys., Austr. 172,1 (text), xi + 188 pp, 2 (plates), 67 Pis. [Repina, L. N.] Penaaa, JL a 1979: Orlow'3\<.I, S, 1959a: Paraoo"loatl 1rom lowtlr M\ooltl Cam­ 3alll!CIDIOCTb },\()'PlloJlol'l!'leCKl!1 brian Strata In the Vicinity of Sandomierz (Central Po­ Upll3HaKOB OT YCJIOBllll OOIlTaHllll TPIlJIot5IlTOB 11 oueinea land). Bull. Acad. polon. Scl., Cl. 3, 7,441-446, 2 Pis. IlX 3HaqeHIl1l ~JIlI CIlCTeMaTIlKll HSJICeMellcTBa 84 Frank Nikolaisen & Gunnar Henningsmoen NGU-BULL.419.1990

Olenelloidea [The dependence of morphological featu­ Sdzuy, K. 1966: Das Kambrium des Frankenwaldes. 2: Die res upon the living conditions of trilobites and an evalu­ Bergeshof-Schichten und ihre Trilobiten-Fauna. Sene­ tion of their importance in the classification of the Super­ kent» leth. 47, 57-86. family OleneJJoideaJ. Akad. Nauk SSSR, Sibir. otdet., Sdzuy, K. 1968: Trilobites del Cambrico Medio de Asturias. Trud. Inst. geol. geofiz. 431, 11-30. Pis. 1-2. Izdat. "Na­ Trab. Geol. " 77·133, 10 Pis. uka", Novosibirsk. [In Russian]. Sdzuy, K. 1978: The Precambrian-Cambrian boundary beds [Repina, L. N.] Penasa, JI.H., (Z. E. PetruninaJ in Morocco (Preliminary Report). Geo/. Mag. 115,83-94, 3. E. neTPYHHHa & (T. I. Khaj rul l inaJ 1 PI. Shergo/d, J.H. 1972: Late Upper Cambrian Trilobites from T. 11. XiUlPYJIHHa 1975: TpHJIOOHTbI the Gola Beds, Western Queensland. Bull. Bur. Min. tr-. lobi tesJ. pp. 100-233, PIs. Resour., geol. geophys. 112 [for 1971], (iv) + 127 pp. 7-48 in Llvanova, Ye. F. 1 llBaHOBa, E.~. ~najdr, M. 1957: 0 novych trilobitech z eeskeho kambria. (ed. ); CTpaTHrpalj)Ha H l\laYHa HHlralerO Vllstnik Ustred. ustav, geol. 32, 235-244, 2 Pis. [In naJIe030a cesepasx npe,l1ropHtl Typl'.eCTaH­ Czech with English summary.] CKoro H AJITatlcKOro xpe6ToB (~bItl ~najdr, M. 1958: Trilobiti iceskeho sttedniho kambria. Rozpr. TaHb-l1BHb) Ustred. ustav, geol. 24, 280 pp, 46 Pis. [In Czech with [Stratigraphy and fauna of Lower Palaeozoic of the English summary.] northern submontane belt of Turkestan and Alai ridges ~najdr, M. 1978: Anomalous carapaces of Bohemian para­ (Southern Tyan-Shan)J. Akad. Nauk SSSR, Sibir. branch, doxid trilobites. Sbomik geol. vea. 20, 7-31, 8 Pis. Minist. geol. UzSSR, Trudy Inst. geo/. geofiz. 278, 352 ~najdr, M. 1984: Revision of the trilobite type material of I. pp. Izdat. Hawle and A.J.C. Corda, 1647. Sbomik nerod. muz. Resser, Ch. E. 1938: Cambrian System (restricted) of the Praze B 39 [for 1983], 129-212, 16 Pis. southern Appalachians. Geo/. Soc. Amer. Spec. Pap. ~najdr, M. 1987: The genera Paradoxides Brongniart and 15, vii + 140 pp,16 pis. Hydrocephalus Barrande (Trilobita). Vllst. Ustfed. !.ist. Richter, R. 1933: Crustacea (Pataontotopls). In Handworter­ geol. 62, pp. 97·104, 2 Pis. buch der Naturwissenschaften. 2nd ed., 2. G. Fischer, [Solovyev, J. A.] COJIOBbeB, 11. A. 1969: Jena, pp. 840-864. (Not seen.) HOB!Ie BH,llbl Paradoxides (TpHJIOCHTbI) Robison, RA 1964: Late Middle Cambrian faunas from H3 rop~~x CJIaHueB aMrHHCKOrO apyca western Utah. J. Paleonto/. 38, 510-566, Pis. 79-92. cesepaoa flKYTHH Robison , RA 1978: Origin, taxonomy, and homeomorphs of Doryagnostus (Cambrian Trilobita). Univ. Kansas [New species of Paradoxides (Trtlobita) from inflammab­ Paleontol. Contr. Pap. 91, 10 pp, 2 Pis. Lawrence, le shales of the Amginskian Stage in northern Ja­ Kansas. kutsk], Nsucnno-issted. Inst. Geol. Arktiki, Minist. Ge­ Robison, RA 1982: Some middle Cambrian agnostoid trilo­ 01. SSSR, Uchen. Zap. (Paleontol. Biostratigr.) 25, 9·20, bites from western North America. J. Paleontol. 56, 4 PIs. Leningrad. [In Russian.) 132-160. stermer, L. 1942: Studies on Trilobite Morphology. Part 11. Rushton, A.W.A. 1966: The Cambrian trilobites trom the The Larval Development, the Segmentation and the Purley Shales of Warwickshire. Palaeontogr. Soc. Lon­ Sutures, and their Bearing on Trilobite Classification. don [Monogr.] 1966, 55 pp, 6 Pis. Nor. Geol. Tidsskr. 21, 49-164, 2 Pis. Rushton, A.w.A. 1979: A review of the Middle Cambrian Strand, T. 1929: The Cambrian beds of the Mj"sen district Agnostida from the Abbey Shales, England. Alcheringa in Norway. Nor. Geol. Tidsskr. 10, 307·365, 2 Pis. 3, 43-61. Strand, T. 1948: On the Middle Cambrian of Hadeland (Os­ Salter, J.w. 1863: On the Discovery of Paradoxides in Brita­ lo Region, southern Norway). Nor. Geol. Tidsskr. 27, in. Q. J. geol. Soc. Lond. 19, 274-277. 89-96. Salter, J.W. 1865: On some New Forms of Olenoid Trilobi­ Stubblefield, C.J. 1936: Cephalic sutures and their bearing tes from the Lowest Fossiliferous Rocks of Wales. on current classification of trilobites. Bioi. Rev. 11, Rept. Brit. Ass. Adv. Sci. 1864, p. 67. London. (Not seen.) 407-440. Salter, J.w. 1869: In Salter, J.W. & Hicks, H. On some [Tchernysheva, N. Ye.] 'lePHbIl:leBaCl H.E. (ed.) Fossils from the "Menevian Group". Q. JI. geol. Soc. 1960: OCHOBbI rraJIeOHTO~OrHH. ~eHHCTO­ Lond. 25, 51-57, Pis. 2-3. HorHe, TPHJIOOHTOOOpaaHbIe H paKOOOpaaHbIe Samsonowicz, J. 1959: On Strenuella and Germaropyge from the Lower Cambrian in the Klimont6w Anticllnori­ [Principles of Palaeontology. Arthropods, trilobito­ morphs and crustaceans]. Gosudarstv. Nauchno­ um. Bull. fAcad. Poton. Sci., Ser. set. Chim., geol. ge­ Teehnich. Izdat. 15, 516 pp. Moscow. [In Russian.] ogr. 7, 525-529, 2 Pis. Samsonowicz, J. 1962 (eds, Korejwo, K. & Teller, L.): Lo­ Thomas, A.T., Owens , R.M. & Rushton, A.W.A. 1984: Trilo­ wer Cambrian fossils from the Klimont6w anticlinorium bites in British stratigraphy. Geol. Soc. London, Spec. of the Holy Cross Mts. (Poland). In Polska Akad. Nauk, Rept. 16, 78 pp. Komitet Geol. Memory book of Professor J. Samsono­ Thoral, M. 1935: Contribution a I'etude pateontoloqlque de wicz. Warszawa, 9-24, 7 Pis. 1'0rdovicien inferieur de la Montagne Noire et Revision Schlotheim, E.F. von, 1823: Nachtrlige zur Petrefaktenkun­ sommaire de la faune cambrienne de la Montagne de. Zweite Abteifung. ? pp, 21 Pis. Gotha. (Not seen.) Noire. TMses Fae. Sci. Univ. Paris A1541 (2407), 362 Sdzuy, K. 1958: Neue Trilobiten aus dem Mittelkambrium pp, 35 Pis. von Spanien. Senck. leth. 39, 235-253. Thcrslund, P. 1935: 11. Paleontologisk-stratigrafisk undersok­ Sdzuy, K. 1961: Teil 11: Trilobiten. In Lotze, F. & Sdzuy, K. ning. In Asklund, B. & Thorslund, P. Fjilllkedjerandens Das Kambrium Spaniens. 1. Abschnitt, pp. 501-595 bergbyggnad I norra Jamtland och Angermanland. Sver. [219-313], Pls.1-15, 2. Abschnitt, pp. 597·693 [315-411], geo/. Unders. C382, Arsb. 28[for 1934}, B6-10B,Pis. 1-2. Pis. 16-34. Akad. Wiss. Lit., Abh. mathem.·naturw. KI. 7-8. NGU-BULL.419,1990 Lowerand Middle Cambrian trilobites 85

Tullberg, SA 1880: om Agnostus-arterna i de kambriska Westergllrd, A.H. 1936: Paradoxides oelandicus beds of aflagringarna vid Andrarum. Sver. geol. Unders. C 42, Oland with the account of a diamond boring through 37 pp, 2 Pis. the Cambrian at Mossberga. Sver. geol. Unders. C '1lidal, G . 1981: lower Cambrian acritarch stratigraphy in 394, Arsb. 3D, 66 pp, 12 Pis. Scandinavia. Geol. Foren. Stockh. Forhandl. 103, Westergllrd, A.H. 1946: Agnostidea of the Middle Cambrian 183-192. of Sweden. sver. geol. unaere. C 477, Arsb. 40, 140 Vidal, G. 1981: Micropalaeontlogy and Biostratigraphy of pp, 16 Pis. the Upper Proterczoic and tower Cambrian Sequence Westergllrd, A.H. 1950: Non-agnostidean trilobites of the in East Finnmark, Northern Norway. Nor. geol. unders. Middle Cambrian of Sweden. 11. Sver. geol. Unders. C 362, 1-53. 511, Arsb. 43 [for 1949], 56 pp, 8 Pis. Vidal, G. 1981: Micropaleontology and biostratigraphy of Westergllrd, A.H. 1953: Non-agnostidean trilobites of the the Lower Cambrian sequence in Scandinavia. In Tay­ Middle Cambrian of Sweden. Ill. Sver. geol. Unders. C tor M.E. (ed.). Short Papers for the Second Internatio­ 526, Arsb. 46 [for 1952], 59 pp. nal Symposium on the Cambrian System 1981. U. S. Whitehouse, F.W. 1936: The Cambrian Faunas of North­ Geol. Surv. Open-File Rept. 81-743, 232-235. Eastern Australia, Part 1: Stratigraphical Outline. Part Vogdes, A.W. 1893: A Classed and Annotated Bibliography 2: Trilobita (Miomera). Mem. oueens: Mus. 11,59-112, of the Paleeozoic Crustacea, 1698-1892, to which is Pis. 8-10. added a catalogue of North American species. Occ. Whitehouse, F.W. 1 939: The Cambrian Faunas of North­ Pap. Calif. Acad. s« 4, 412 pp. San Francisco. (Not Eastern Australia. Part 3: The Polymerid Trilobites (With seen.) Supplement No.l). Mem. Queensl. Mus. 11, 179-282, Vogt, Th. 1967: Fjellkjedestudier i den 0stlige del av Troms. Pis. 19-25. Nor. geol. unoers. 248, 1-60, 2 Pis. Whittington, H.B. 1988: Hypostomes and ventral cephalic Wrern, B. 1952: Palaeontology and Stratigraphy of the sutures in Cambrian trilobites. Palaeontology, 31, Cambrian and Lowermost Ordovician of the BOdahamn 577-609, PI. 52-55. Core. Bull. Geol. Inst. Univ. Uppsala 34, 223-250, 1 PI. Whittington, H.B. & Kelly, S.R.A. 1983a: Morphological terms Wahlenberg, G. 1818: Petrificata Telluris Svecanae. Nova applied to Trilobita. 29 pp. Written communication Ap­ Acta R. Soc. Sci. Upsala 8 [for 1821], 1-116, 4 Pis. ril, 1983. Upsaliae. (Not seen. According to bibliographical note Whittington, H.B. & Kelly, S.R.A., 1983b: Morphological terms by Bassett & Cocks 1974, p. 40, the main body of this applied to Trilobita. Supplement to revision of April work was published and distributed in advance of the 1983.9 pp, 7 Pis. Written communication August, 1983. serial issue.) Yang Yia-Iu 1978: [Middle and Upper Cambrian Trilobites Walcott, C. 1910: Cambrian Geology and Paleontology. No. of Western Hunan and Eastern Guizhou]. Chin. Acad. 6. Olenellus and other genera of the Mesonacidre. Geol. Sci., Pr of. Pap. Stratigr. Palaeontol. 4, 1-82, Pis. Smithson. Mise. Coli.53, 231-446, Pis. 23-44. 1-13. Geological Publishing House, Peking (Beijing). [In Wallerius, I.D. 1930: Frlln VlistergOtlands mellankambrium. Chinese with English abstract.] Geol. Foren. Stockh. Forhandl. 52, 47-62. Yang Yia-Iu 1982: [Notes on the Middle Cambrian trilobite Warburg, E. 1925: The Trilobites of the Leptrena Limestone faunas from Duibian of Jiangshan, Zhejiang]. Geol. in Dalarne. With a Discussion of the zoological Position Review 28,299-307, 2 Pis. [In Chinese with English and the Classification of the Trilobita. Bull. Geol. Inst. abstract.] Univ. Upsala 17, vi + 1-446, 11 Pis. Yin Gong-zheng & Li Shan-ji 1978: (Trilobites]. In The Welsch, M. 1983: A newly discovered acritarch sequence Working Group of Stratigraphy and Palaeontology, from Middle cembrtan to Tremadoc continental margin Guizhou section (ed.). [A Handbook of Palaeontology deposits of the Digermul Peninsula, Finnmark, Northern ofsouthwest China. Guizhou Part (1)),385-594, 798-829, Norway. Palynology 7, 254-255. Pis. 144-192. Geological Press, Beijing. [In Chinese.] Welsch, M. 1984: Die stratlgraphische Bedeutung einiger Zheu Tlen-mal, Uu Yi-iang, Meng Slan-sung & Sun Zhsng­ ausgewlihlter Acritarchen aus Sedimenten der Diger­ hua 1977: (Trilobita]. In Wang Xiao-feng & Jin Yu-qin mul-Gruppe (Mittelkambrium bis Tremadoc), Finnmark, (eds.). [Palaeontological Handbook of Central and Sout­ Nordnorwegen. Geol. & Palaeontol. 18, 5-7. hern China. Central-southern area palaeontology. Part Welsch, M. 1986: Die Acritarchen der Mheren Digermulgrup­ 1. Lower Palaeozoic Section), 104-266, Pis. 36-81. pe, Mittelkambrium bis Tremadoc, Ost-Finnmark, Nord­ Earth Science Press, Beijing. [In Chinese.] Norwegen. Palaeontographica Abt. B, 201,109 pp, 9 Pis. Zenker, J.C. 1833: Beitrage zur Naturgeschichte aer ur­ Westergllrd, A.H. 1930. In Holm, G. & Westergllrd, A.H. A welt. Organisch~ Reste (Petrefacten) aus der Alten­ Middle Cambrian fauna from Bennett Island. Zap. Nauk burger Braunkohlen-Formation, dem Blankenburger Soyuza SSR, Ser. 8, Otde/. fiz.-matem. nauk 21 (8), Quadersandstein, Jensiscben bunten Sandstein und 25 pp, 4 Pis. Leningrad. Bohmischen Uebergangsgebirge. viii + 67 pp, 6 Pis. Jena. (Not seen.)

Manuscript received October 1989; revised typescript accepted March 1990. 86 Frank Nikolaisen &Gunnar Henningsmoen NGU-BULL419,1990

Fig. 6. All specimens figured in dorsal view. A-G. Peronopsis terox (Tullberg, 1880) sal/esi (Bergeron, 1889). Lower trilobite horizon in the middle Middle Cambrian sandstone and shale member (K2) of the Kistedal Formation. Locali­ ty 12. Collection M.-L. Windolph & M. Welsch1981. A. Fairly complete but slightly distorted and exfoliated cephalon. PMO 111.589. x 9. B. Latex cast of external mould of incomplete but only slightly distorted cephalon. PMO 111.593. x 10. C. Small but undistorted cephalon. PMO 111.603. x 18. D. Nearly complete and very slightly, transversely compressed internal mould of pygidium. PMO 111.599. x 12. E. Latex cast of external mould of exfoliated pygidium with second tergite attached. PMO 111.598. x 10. F. Latex cast of external mould of incomplete but only slightly distorted and flattened pygidi­ um. PMO 111.588. x 9. G. Latex cast of external mould of slightly crushed pygidium with very poorly preserved tho­ rax attached. PMO 111.600. x 9.

H-M. Doryagnostus incertus (Bn.~gger, 1878). Middle Middle Cambrian sandstone and shale member (K2) of the Kistedal Formation. H. Fragmentary cephalon. PMO 86437. Locality 7. Collection H.G. Reading 1959. x 9. I. Fairly complete but sagittally compressed pygidium. PMO 72382. Locality 7. Collection as for H. x 9. J. Latex cast of external mould of small pygidium with attached but poorly preserved thorax and posterior part of cephalon. PMO 86292. Locality 10. Collection as for H. x 12. K. Latex cast of external mould of fragmentary pygidium. PMO 72378. Locality 7. Collection as for H. x 8. L. Large, almost complete but poorly preserved internal mould of pygidium. PMO 111.605. Upper trilobite horizon at locality 12. Collection M.-L. Windolph & M. Welsch 1981. x 7. M. Latex cast of external mould of nearly complete but slightly distorted pygidium. PMO 86480. Locality 11. Collection as for H.x 7. NGU • BULL. 419, 1990 Lowerand Middle Cambrian trilobites 87 88 Frank Nikolaisen & Gunnar Henningsmoen NGU- BULL. 419.1 990

Fig. 7. A-C. Kjerulfia lata Kicer, 1917. Lower Cambrian massive-bedded quartzite member (02) of the Doulbasgaissa Formation. Scree at the outcrops on the northwestern slope of Bardeviktind. A. Dorsal view of fragmenta ry specimen show ing right half of genicranium lacking palpebral lobe. PMO 82712. Original of Henningsmoen 1984, p. 23, fig. A and Ahlberg & Bergst rom 1986, Figs. 1, 8. Collection G. Henningsmoen 1963. x 1 1/2. B. Dorsal view of latex cast of very fragmentary genicranium. PMO 98551. Collection K.E. Henningsmoen 1963. x 1 1/2. C. Dorsal view of latex cast of fragmentary glabella. PMO 98549. Collection F.Nikolaisen 1963. x 1 1/2.

0-1. Ellipsocephalus et. hoffii (Schlotheim, 1823). Middle Cambrian quartz ite and shale mem­ ber (K1) of the Kistedal Format ion. Northeastern slope in Kistedal (Gistovagge). D. Dorsal view of fragmentary anterior part of incomplete axial shield. PMO 72350. Collection G. Henningsmoen & F. Nikolaisen 1960. x 2. E. Dorsal view of fairly complete but internal mould of cranidium. PMO 72355. Collection as for D. x 4. F. Dorsal view of fragmentary axial shield. Anterior part of specimen figured is a cast made of silicon rubber from the counterpiece. PMO 72348 and 72349. Collection H.G. Reading 1959 x 2 G. Dorsal view of internal mould of fragme ntary cranidium. PMO 72356. Collection as for D. x 2. H. Dorsal view of internal mould of fragmentary cranidium. PMO 72354. Collection as for D. x 2. I. Dorsal view of internal mould of fragmentary cranidium. PMO 72345. Collection as for F. x 2. NGU -BULL419. 1990 Lowerand Middle Cambrian trilobites 89 90 Frank Nikolaisen & Gunnar Henningsmoen NGU - BULL. 419. 1990

Fig. 8. A-E. Hydrocephalus ct. carens Barrande, 1846. Lower Middle Cambrian quartzite and shale member (K1) of the Kistedal Formation. A. Dorsal view of fragmenta ry cranidium. PMO 72403. Locality 4. Collection F. Nikolaisen 1960. x 1. B. Dorso-Iateral view of latex cast of external mould of almost comp lete right librigena. PMO 86231. Locality 4. Collection G. Henningsmoen & F. Nikolaisen 1960. x 2. C. Ventral view of incomplete hypostome. PMO 86235. Locality 4. Collection as for B. x 3 1/2. D. Dorsal view of nearly complete right thorac ic pleura. PMO 86231. Locality 4. Collection as for B. x 2. E. Dorsal view of very incomplete and very poorly preserved thorax. PMO 86584. Locality 3. Collection M.F. Tuke 1961. x 1 1/2.

F. Eccaparadoxides ct. pusillus (Barrande, 1846). Dorsal view of fragmentary and very poor­ ly preserved cranidium and two axial rings. PMO 86583. Most probably lowermost part of the lower Middle Cambrian quartzite and shale member (K1) of the Kistedal Formation. Locality 2. Collection M.F. Tuke 1961. x 3 1/2.

G-1. Paradoxides davidis Salter, 1863. Middle Middle Cambrian sandsto ne and shale member (K2) of the Kistedal Formation. G. Dorso-Iateral view of almost complete left librigena. PMO 72390. Locality 10. Collection H.G. Reading et al. 1959. x 2. H. Dorsal view of small and slightly incomplete dorsal exos keleton. PMO 111.608. "Upper trilobite sequence" at locality 12. Collection M.-L. Windolph & M. Welsch 1981. x 6. I. Dorsal view of latex cast of the external mould of the dorsal exoskeleton in H. PMO 111.609. Same locality and collectors. x 6. NGU - BUL L. 419, 1990 Lowerand Middle Cambrian trilob ites 91 92 Frank Nikola isen & Gunnar Henningsmoen NGU - BULL. 419, 1990

Fig.9. A-G. Paradoxides davidis Salter, 1863. Middle Middle Cambrian sandstone and shale mem­ ber (K2) of the Kistedal Formation. A. Dorsal view of almost complete but fairly small and sagittally slight ly compressed cranidi­ um. PMO 111.596.'Lower trilo bite sequence ' at locality 12. Collection M.-L.Windolph & M. Welsch 1981. x 6. B. Dorsal view of latex cast of external mould of almost comp lete but small cranidium . Note difference in size between the two palpebral lobes. PMO 111.609.Locality and collectors as in A. x 6. C. Ventral view of almost complete but slightly sagittally compressed rostri-hypostomal pla­ te. PMO 111.586. Locality and collectors as in A. x 6. D. Dorsal view of fragmentary and slightly distorted cranidium. PMO 111.601. Locality and collectors as in A. x 6. E. Dorsal view of incomplete and strong ly flattened cranidium. PMO 72391. Locality and collectors as in E. x 2. F. Dorsal view of incomplete cephalothorax. PMO 72396. Locality 10. Collection H.G. Reading et al. 1959. x 1. G. Ventral view of slightly incomplete and somewhat distorted rostri-hypostomal plate. PMO 72379. Locality 7. Collection H.G. Reading et al. 1959. x 5. NGU - BULL . 419, 1990 Lower and Middle Cambrian trilobites 93 94 Frank Nikolaisen & Gunnar Henningsmoen NGU- BULL. 419.1990

Fig. 10. A-F. Paradoxides davidis Salter , 1863. Middle Middle Cambrian sandstone and shale mem­ ber (K2) of the Kistedal Formation. A. Dorsal view of small and slightly distorted but almost comp lete cranidium. PMO 111.587. 'Lower trilobite sequence ' at locality 12. Collect ion M.-L. Windolph & M. Welsch 1981. x 81 /2. B. Dorsal view of incomplete and distorted cranidium showing long postocular facial suture. PMO 72445. Locality 16. Collection G. Henningsmoen 1960. x 1 1/2. C. Dorsal view of latex cast of five thoracic tergites from the posterior part of the thorax . PMO 72393. Locality 10. Collection H.G. Reading 1959 et al. 1959. x 4. D. Dorsal view of almost complete cranidium and anterior part of thorax, most probably repre­ senting a moult assemb lage. PMO 86283. Locality 7. Collection as for C. x 2. E. Dorsal view of latex cast of large pygidium. PMO 72394. Locality and collection as for C. x 3. F. Dorsal view of latex cast of externa l mould of complete but slightly sagittally compressed pygidium with ankylosed incomplete thoracic terg ite. PMO 111.602. Locality and collectors as for A. x 5. G. Ptychopariine gen. & sp. indet. Dorsal view of latex cast of external mould of small and poor ly prese rved cranidium. PMO 72453. Middle Middle Cambrian sandsto ne and shale member (K2) of the Kistedal Formation. Locality 16. Collection G. Henningsmoen 1960. x 6.

H-J. Nassovia? mjol/nir n. sp. Middle Middle Cambrian sandstone and shale member of the Kisteda l Formation. H. Dorsal view of paratype left, incomplete librigena show ing long anterior doublure. PMO 72389. Locality 13. Collection H.G. Reading et al. 1959. x 2. I. Dorsal view of fragmentary pygidium most probably belong ing to this species. PMO 86143. Locality 16. Collection F. Nikolaisen 1963. x 4. J. Dorsal view of holotype, flattened but almost complete cephalon with anterior part of the thorax attached . PMO 72388. Locality and collectors as for H. x 2.

K-L. Anomocarina? sp. Middle Middle Cambr ian sandstone and shale member (K2) of the Kistedal Formation. K. Dorsal view of latex cast of incomplete and poorly preserved pygidium. PMO 72375. Loo­ se block at locality 17. Collection G. Henningsmoen 1960. x 2. L. Dorsal view of latex cast of externa l mould of small fragment of pygidium. PMO 72371. Locality 14. Collection H.G. Reading et al. 1959. x 1 1/2. NGU · BULL. 419,1990 Lowerand Middle Cambrian trilobites 95