OLENELLUS (TRILOBITA) FROM THE LOWER CAMBRIAN STRATA OF NORTH-WEST SCOTLAND by JOHN Cow1E and KENNETH J, MCNAMARA Atsnacr New ater of Olen pmo Pech Hore, 192.0, Peach 194,240. ermets en enables edption a hese orm ad ustrson by popper he fist ne tas Pach, #84 suppres and 0. hamocss.nv. serted. Grape a se toilet he cations ena ‘Tho fauna represents the Yount pr he Boone -Olen Zone te Pai Prone oe Oolsd Rea Tae Cambro-Ordovician outcrop of north-west Scotland is about 200 kilometres long and varies in width betwcen 2 and 20 kilometres stretching from Duress in the north to Skye. The Cambro-Ordovician beds were evidently laid down in a sea which gradually transgressed across older, Precambrian, strata, resting with angular un- conformity on them. Total thicknesses are not well known, partly due to the nature of the outcrops nd the involvement of the strata in a major thrust-zone. The olenellid- ‘bearing formations are the “Fucoid’ Beds (12-27 m thick) and the overlying Salteella Grit (5-15 m thick), ‘The *Fucoid’ Beds are dolomitic siltstone or shale, arenaceous dolomites, arena- ceous dolomitic mudstones, and grits with ripple-indueed micro-cross-laminations, in places thin limestones and lenticular dolomitic sandstones occur; the weathering is characteristically brown to buff and ferruginous in colour, Swett (1969, p. 645) ‘considered that the “Fucoid” Beds were deposited in shallow to moderately deep water on an ever-broadening shelf at increasing distances from the detrital sources, Thin series of beds of grey siltstone, arenaceous shale or shale, which weather rusty-brown, are the main source of olenelid trilobite remains which are generally isarticulated exuviae, though occasional articulated specimens have been found. ‘The specimens are generally preserved as internal and external moulds. Due to the very thin nature of the exoskeleton there are no significant differences between internal and external moulds, apart from the presence of the ornament on the external ‘mould, though in some instances it has been impressed through to the internal mould, (Pl. 70, fig. 12). Fossiliferous horizons occur at several levels in the “Fucoid” Beds (Brand 1965). ‘The Saiterella Grit is a thin persistent grit formation predominantly made up of quartzite, orthoquartzitic sandstone containing minor detrital carbonate grains, arkosic grit, dolomitic sandstone and sandstone, but there are interleavings of shales near the base which give Olenellus lapworthi. The fossil collections are meagre com pared with those from the ‘Fucoid’ Beds. ‘The first specimens of trilobites from Scotland assignable to Olenellus werecollected in the late nineteenth century by M. Macconochie, a member of the Geological Survey team led by B. N. Peach and his colleagues. The trilobites came from Allt (Pata, Yo 2 Fe 1 S64 6-7as PALAEONTOLOGY, VOLUME 21 ‘nan Righreon (Allt Righ Tan) in the Dundonnell Forest, Ross and Cromarty (Peach and Horne 1892, p. 228) (text-fig. 1). Around the turn of the century further dis- coveries of olenellid trilobites were made along the north-west Scotland outerop strip of the Fucoid Beds and Salterella Grit (see text-lig. 1). Localities mentioned in the text and shown on textfig. 1 are An-t-Sron, Loch Eriboll (1), Skiag Bridge (2), Loch Awe quarry (3), Knockan Cliff (4), Ailt nan Righteon and Loch an Nid (5), Meall a’Ghiubhais, Kinlochewe (6),and Ord, Skye (7). In the 1960s and 1970s new collections of olenellid trilobites were made by the Institute of Geological Sciences (Bowie et al. 1966; Brand 1965) and by John Cowie, Kenneth McNamara, Susan Radford, and John Branegan from various localities including especially the recently excavated roadstone quarry at Loch Awe, south of Inchnadamph, Ross and Cromarty (see above and text-igs. 1-3). Descriptions and illustrations of some of the fossils dealt with in this paper were published by Peach ‘and Horne (1892), Peach (1894), Peach er al (1907), Walcot (1910), and Lake (1937) but in these earlier works only drawings were used without photographs. ‘The purpose of this paper is to redescribe the species in the light of the new dis- ccoveries at Loch Awe, and to describe a new species, Olenellus hamoculus. Specimens were qualitatively designated; significant variable features were then measured and plotted on graphs (text-figs.2,3)to illustrate the relationships established. Deserve terminology and igus. Terms used inthe systematic desrption follow Haingon ¢! a (or Moore 1959) exept that the terms ‘extraocular are and inte-ceuar te are wed in place "ena (Moore 1959, p. O10); “imtergsnal spine’ is preferred to metagenalspine’- One new tr, “pialpebal Tuzrow, introduced to describe the lngitainal furrow i the ee lobe “The parameters ploted in tex-g. 2 and 3 are associated with those morphologic atures which re ‘often described s being of specif significance among olenlids, eg relative kngth of eye lobe (ext Hg, 2, by, label shige (et. 30) relative length of preglabelr fil ex-f ) "Teas shows thatthe relative sagital length ofthe cephln and preslaelia eld has ile peciic ‘importance whe diferentiating between thers thre olenlige, Text-fg.2vand lustrate the importance ‘fey length, whit tex a shows how the gabe ape can vary interopecieal ‘The numberof speximens measured, m, varies due vo some cephsla being incomplete 40 that not all parameters were obtain ‘All measurements were made with eye lobes 4 horizontal plane, Associated faunas and the zone represented In addition the fauna includes: the trilobite Olenelioides armatus, Peach; the brachiopods: Micromitra sp., Lingulella zeus Waleott, Paterina sp. Kutorgina sp. Aerothele subsidua White, “Murchisonia? sp.; miscellanea: Hyolithes Sp, Saltereltaspp., Coleoloidessp.,echinodermata ind, rustaceaind,,and the worm trace fossil Planolites sp. Elements ofthe faunal assemblage probably correlate with those from the younger pact ofthe Lower Cambrian given in Fritz’ (1972) Bonnia-Olenellus Zone, although the Scottish trilobite fauna is limited in ts representation of genera. Fritz (1972, p.7) lists genera that are commonly present inthe zone as Ariagmus, Bonnia, Laudonia, Olenelus, Onchocephalus, and Zacanthapsis. The base ofthe Bonnia-Olenelus Zone is placed by Fritz atthe fist appearance of Olenelas. The underlying Nevadelfa Zone contains the diagnostic genera Bradyfallowspis, Holmiella, and Nevadella but Olenetus is, by definition, absent. Olenellus is also absent in strata overlying the Bonnia-Olenellus Zone‘exr-ig |Past of aorth-west Scotland with selestadFosslfrous Lower Cambrian ‘Facid’ Beds localities (National Gri erences sven) along the outrop (chown diagrammatically by a black ine from Dumnss to Skye, An--Sron, Lax Enbl (NC #42 82): 2 Shag Bridge (NC 288 249) 5, Loch Awe quarry (NC 251 158) 44 Knockan Cli(NC 18209): 5, Allan Righton (NH1O3S 194) and Loch an Nid (HPORS 747); 6, Meall @Ghitbhas, Kinlochowe (NH 979 685), 7, Ocd, Skye (NG 613 133),sail itnc rom oui of ee be fo pwr of cate PALAEONTOLOGY, VOLUME 21 +0, lapwerthi; 0.30 ° #20 hamocuiue, 22 k= O.retieuiatues 88 - « (a e . 20. lapworthi, 0.27 ee #0 hameculus; 1 25 a 150, reticwlatus: 9.31 xo 2) sexi 2a, difrentiaton of 0. lapworh fom 0. reticulata and 0. hanes2+ Qlopworthis +O homocuius; 0 reticulate n.26 1.25. noe 3 7 vo 7 2+ 0. lapworthi: 9.27 : 8: Ohemoculus; 0. 24 4+ Oreticulatus: 0. 60 7 Soa eng of cepa) oy oy ‘TXT. 2adifeentition of 0. hamocdafrom O. lapwordhisnd O. eins, reation of peal "el to sapial cephalic lengthoo PALAEONTOLOGY, VOLUME 21 Age equivalence between the ‘Fucoid’ Beds of Scotland and the Bastion Formation of East Greenland (Cowie and Adams 1957) can be based on the faunas which include fairly closely comparable species of Olenellus from similar lithologies. Spocimens are lodged in the Institute of Geological Ssiences, in Edinburgh (GSE) and in London (OSM), fhe Bes scum Naira Hy (BM), ad the Depart of Gzlogy, Unie of Sc ‘Andrews SYSTEMATIC PALAEONTOLOGY Family oLeNtLupar Vogdes, 1893 ‘Gens OLENELLUS Billings, 1861 Type species. Oloms thompson Hall, 1859, from the Lower Cambria of Geoesia Township, Vermont Remarks, The relationship between Olenellus and Paedeumias has been discussed recently by Fritz (1972) and Cowie (1968). For the purposes ofthis paper the alloca tion of Paedeumias to Olenellus suggested by Fritz is accepted. Oleneltus lapworthi Peach and Horne, 1892 Plat, 1-6 Ooms lpworth Peach and Home, pp. 227-242, pl. 5, ‘Ona lapworthi Peach and Hore, . figs. 79 101,13, non ‘Ons igor Peach; Peach, pp. 662-66, p29, Olnais apworth vat longa Peach, p. 6, 2, ig. 4 (enc igpwort Peach and Hore; Peach ea. p. 401, 04, 1 (Oona gpword Peach; Walet, pp 381-332, 3, is. 46, (Ofna lapworthi Peach: Lake, pp. 238-240, pl. 33, fig 10, ow IL p34, fg. 1 2,5 mh EXPLANATION oF PLATE 69 Allspesimens from the Paco Bes Figs. 1-6. Olena lpworsi Peach and Hore, 18721, GSE S364, ntenal mould of lctotype ephalon from Altnan Rigteon, Dundonpel: «1's Figured by Peachand Hore 1892 ps, 5-2. 2, GSES300, late east of external mould of incomplete dorsal exoskelston from Mall Giubhals, Kinlochewe: “8. Figured by Walt 1910 p39, fg 5. 2-5, OSE 1330, 13306, 13310, terol movie of ephala fom Loch Awe quarry, near Inchaadamph: =, 2, 25! 6, BMI 1118, sternal mould of fag ‘mentary dorsal exoatleton showing impression of hypostome om slab rom Meal "Guth, Kinlosheve: 2 Figs 7-15 Ole resicuars Peach, 1894. 1, GSE S343, ineral mould of leteype cephaton from ‘sm locality as Fig 6; > 15.8, BMI 1110, internal mould of hypostome fom sume local as Fig. 6: ~ 3.9, GSE S34, intern mould of paralecorype cephalo fom Alt nan Riphreon, Dundoael: Figured by Peach 1854, pl 30. 3-10-12, GSE 1829s, intemal mould of eephulon fom Loch Awe ‘quarry near Inchadap, Doral fv-lateal, and anterior views: 2-18.14, OSE 13324, 13335, intemal moulds of small cephala from sue locality a8 is. 10-12, <4, 3.” 1S, OSE $342, eternal ‘mould of paralectotypecepealon from Mall wGhiubhas, Kilocheve! ~ I. Figured by Peach 198, v0, 2PLATE 69 ‘COWIE and MCNAMARA, Olenellusoa PALAEONTOLOGY, VOLUME 21 1937 Wamera? sp: Lake, pp. 246-247, pl. 35, fg. 5 1951 Padeumiashanceni Poulsen: Poulsen, p. 16 11957 Olena lpwors Poach; Passer, p. 120. 1960, Olenelusapworeht Peach: Kila, p37 1865 Olenelus apworcht Peach: Brand. 286 1972. Olenetus apworthi Peach; Covi a. . 2 1972. Paedeumishansent Poulsen: Cowie ea p28. 1974. Olenetisapwortht Beach: Cows, . 136. 1974. Paedewmias hansen! Poulsen; Cowie, p16. Lectotype. An incomplete cranium, GSE $368 (PI 9,1), fom the "Fucoid’ Bos, Lower Cambrian at Altman Righteon (All Righ Lan) NH 095 798 near the hllirsck7 kn south of Dundonnel, Ros and ‘Cromarty, Sctind: igured by each and Horne (892, p52); hee selected, Dimension of lecorpe. wat Sagittal length of cephalon 13 ‘Maximum with of cepa a4 Sagital dimension of preglabelar ld 14 etal length fom anterior cephalic margin to anterior glabellar furrow Jp) «93, Exsagital length of ee lobe ” Exsapital distance from posterior Up of je lobe to posterior border of cephalon 25 Paralecotypes Incomplete cephala GSE 458, GSE 457, GSE 456, GSE 536, gut by Peach and Horne (92, ples 3-6 respectively); same horizon and locality as stonype, Morera rion and cas Peach and Horae based this pies. ona smallamount of poorly preserve, fragmentary materia rom All nan Righteon which was figured in thir paper of 189, ln 189 collections {Hom Meall aGhubhais gave bette evidence. Though a le Mateed in sale he fos are quite well ‘reser and contin some complete specimens. opworthcecurs ite moe frequenty atti caliy {han 0-reticadausPeach, 1898, Recent collections rom he quarry a¢ Loch Awebaveyteied approximately twenty tvewell preserved exphals of O lap ih and numetonsagments; here thisspecesisinarninoriy, comprising about 17, othe otal umber of wel preserved oleneliécepala. Thorac segment ar 7 ead af lost invariably found lated hae making the speci tsgnrent itil Diagnosis, Preglabellar field long; occupies one-fifth cephalic length in small speci- ‘mens, one-tenth in long ones; frontal lobe well-rounded anteriorly; eye lobes are strongly curved and long, reaching at least as far back as the oceipital ring; small inter-ocular area. Posterior border almost straight with genal angle in transverse line with occipital ring. Posterior of frontal lobe less than half cephalic length (sag.) from posterior of occipital ring. Exoskeleton faintly reticulated Description. Cephalon varies between semicircular (Pl. 69, fig. 3) and paraboloid (P1.69, fig. 4), the majority being semicircular; bordered by narrow, convex rim that slightly widens and flattens as it approaches genal angle. Posterior border very narrow at axial furrow, widening to genal angle; directed transversely out from axial furrow initially, then curving slightly forward distally to slightly advanced genal angle. Intergenal angle two-thirds posterior margin length from axial furrow; border thickens slightly at intergenal angle (Pl. 69, fig. 4). In immature specimens intergenal angle at half posterior width from axial furrow and genal angle in a more advanced position (PI. 69, fig. 2). Genal spine continues line of lateral border, being directed slightly outward posteriorly; extends for over half length of cephalon. Glabella ‘widest anteriorly narrowing to 1p furrow then slightly widening to occipital ring. Frontal lobe moderately convex and occupies more than half of glabellar length inlarge specimens, les than half install ones; occupies one-quarter of cephalic widths well rounded frontally (Pl. 69 fig. 3); posterior stuated at over half cephalic length from anterior border. Ratio of cephalon length to preglabellar and frontal lobe length varies between 1:0:53 and 1:05. The frontal lobe is almost equal to the width of the occipital ring. In 0. reticulatus the Trontal lobe ia litle broader relatively. There are three pais of glabellar furrows: 3p furrow transplabellar directed strongly forward abaxilly at 45", then curves back through aright angle to direct backward as it enters axial furrow, 2p furrow not transplabllar: parallel to anterior furrow adanially, but abaxially becomes divergent resulting in anterior glabellar lobe lengthening (exsag) distally, nearly geniulate then narrowing abruptly close to axial furrow; gently recurved backward at axial furrow where it abruptly shallows (PI. 6, fig. 5) Sp and 2p Furrows situated close to each other, resulting in 3p lobe being narrower (sag. and exsag.) than 2p lobe as Ip furrow is situated further from 2p furrow. Ip furrow not transplabellar: wider (exsag.) than anterior and median furrows, consstingofshort, abaxial situated slits directed gently forward abaxial) Occipital furrow similar to posterior plabellar furrow. Node situated postero-medialy on occipital ring (PI. 69, fig. 1). Moderately long preglabellar field (PI. 69, fig. 3) ‘occupies one-tenth or more label length in large specimens, up to one-fifth n smal ones (PL 63, ig. 2, From postero-lateral part of frontal lobe eye lobe arches out Strongly, making an angle of about 140° with frontal lobe. It is Tong (lext-fs. 20), ‘ceupying two-ihs cephalic length in mature specimens, one-half in small ones, Strongly curved, wide (¢r) with gently convex dorsal surface; wellcurved inner ede, tore strongly curved outer edge, eye lobe being furthest from glabella opposite Ip furrow; posteriorly terminates opposite and close to occipital rng, in some examples (P16, 3) almost reaching posterior border; postoctlar cephalic length is about ‘one-tenth total cephalic length (text-fig. 2a). Eye surface occupied outer edge of eye lobe, being now represented by a narrow furrow (PI. 69, fig. 5). Shallow epipalpebral furrow is sometimes present. Inter-ocular area almost enclosed; small and slightly raised. Extra-ocular area broad, low, sloping very gently laterally. Hypostome is tinknown. Thorax composed of fourteen segments (P. 69, fig. 6); third segment macro-pleural, being longer (exsag.) than any of other pleurae, and extended postero- laterally as long pleural spine. Axis tapers gradually posteriorly. In mature specimens (PI. 69, fig. 6) pleurae (to forum) occupies slightly more than one-third thoracic ‘with anterioryas they narrow (i) posteriorly more strongly than axis. In immature Specimens pleurse occupy less than one-third thoracic width anteriorly. First, scond, fourth, ith pleurae develop very short spines laterally those posterior to fithsea2 ment develop longer pleural spines. leurae depressed medially, bounded anteriorly and posteriorly by rased pleural bands, Posterior axial ring bears long spine which iis almost as long as thorax (PI. 69, fig. 6). Discussion, Peach’s paper (1894), in which further species of Olenellus were described, iustrated specimens, many of them arcuate, trom the Fucoid Beds of Meal a’Ghiubhais, Kinlochewe. As suggested by Peach (1894) the specimens of O. lapworthi from Mealls'Ghiubhais are generally smaller than the O. revclatue specimens at the converse is true at the Loch Awe locality. This species shows small depres of intraspecific variation, a has bon alluded (0 in te description; this involves smallon PALAEONTOLOGY, VOLUME 21 variation in length of preglabellar field (contrast PI. 69, fg. 3 and PI. 69, fg. 6), eve lobe (contrast Pl. 69, fig. 3 and Pl. 69, fig. 1, position of genal angle, and over-all cephalic shape (contrast Pi. 69, fig. ! and Pl. 69, fig. 4). The specimens described by Peach and by Peach and Horne tend to be semicircular, but many collected from Loch Awe tend toward a more parabolic outline. Comparisons with other forms present in the Fucoid Beds will be made under the relevant discussions. Other species similarto 0. lapworthiare: O.transitans(Walcott, 1910), 0. thompsoni (Hall, 1859), 0. hanseni (Poulsen, 1932), and 0. svalbardensis Kielan, 1960). 0. transi- ‘ans can be distinguished by its narrower glabella and slighty longer eye lobe, more distally positioned intergenal angle, larger inter-ocular area and smaller frontal lobe (see Walcott 1910, p. 38), whilst O. thompson, another North American form, lacks a preglabella field and possesses a broader border. O. hanseni, from the Lower ‘Cambrian of Ella Island, East Greenland, is very similar to O. lapworthi,diflering only in having a slightly longer eye lobe which reaches to the posterior border. (0. svalbardensis from the Slakli Series in Spitsbergen, possesses a shorter preglabellar field, broader and more advanced genal angle. Olenettus reticulatus Peach, 1894 Plate 6, is. 7-15; Pate 7 fs, 12,12 1892 Olen apworii Peach and Horse, pl 5, fs. , 7,8, 1834 Oleelretearas Peach pp. 663-426, i 30, fas. 1-3, 76-14; p31 is. 17. 1894 Oleel lao each pas): Peach, pl. 29,2, 5 1894 Oleel lgpworh va. elomgaras Peach (pars), p29, 3. 1894 Olenell gar Peach, cx l .667 1907 Oleel reevans Peach; Peace ap... 4, 4,6 1907 Olenelis gpworat Peach: Peach ea, 82 is. 2,3 1910 Olea retain Pech; Was p33 3 pL 3S 1,714 1910. Olenlies epwors Peach (pars; Wale, 39, Bit. Olek elans Pal ake py 380208, fs 79 10,11 1939. Ole lapwor Poach (par) Lake, p34 is 1, 3046 ‘1937 WameriaTep.; Lake, pp. 246-247 (pars), pl.35,08.4. 1951 Wanreri nathorai Poulsen; Poulsen. i 1957 Olenelis eicolaru Peach: Palmer, p. 120 1972. Olenelts eicularus Pach: Conte ap 28. 1912 Wanner maths Poulsen Cowie ot ap 28. 1914. Olenihr eieutana Peach; Comte. 13 1974 Wanner nathrat Poulsen Covi, p 136 “ectonype. An incomplete cepalon, GSE 5343 (PL 69,7) from the “Fucoit’ Beds, Lower Cambrian on te northern ope of Meal « Ghiubhas, 5 km WNW. of Kinloenewe, Ross and Cromarty, Sot Figured by Peach (1894, p30, 1); herein elected Dimension of lectorpe 255 (estimates) 20 estimated) 2 Sogial eg fom enero cepa margin to aro lar Turrow (3p) as Esai length ofee lobe 85 Exagtal distance fom posterior ip of ee lobe to posterior border or ephalon =Paralectorype. Two incomplete exphala, GSE 532 and GSE 5344, on incomplete articulate specimen, {GSE 550, and an ncompet thorax GSE 5351; same Horizon and locality as lcotype: hgued By Peach (1894, pl. 0, ps 2.4 spl, Bp 4 respectively), Matera orzo and ocean ote muerous, wel preserved hough generally incomplete, ‘expats from the type locality at Meall x Ghiubbais, ths species has been found at All nan Righreou (Peach and Home, 1992, pl fig. 1; this specimen has ashore exe lobe tan is evident from Pooch and Horne drawing). Recent collcing at the Loch Awe quarry (ext 1) has 100 wel expla snd countess fragmentary specimens. Ths specion occurs le es frequent 'Ghiubhais than Oapword, but at the Lach Awe quatty itcomprises about 65% ofthe elenel [Atal these localities itis ested {0 the Fuco Bas, Lower Cambria Diagnosis. Preglabellar field short; frontal lobe tapered anteriorly. Eye lobes short, posterior tips being in line with Ip lobe and gently curved. Genal angle slightly ‘Advanced from normal position, distal part of posterior border is concave forward from intergenal to genal angle. Exoskeleton coarsely reticulate. Remarks. This species differs from 0. lapwortht in the following respects: (@ preglabellar field shorter, occupying one-tenth cephalic length in small indi- ‘viduals, and more than one-tenth in large ones; (©) frontal lobe tapers anteriorly (Pl. 69, figs. 14, 15); posterior set at one-half cephalic length from anterior margin: (©) frontal lobe occupies one-third cephalic width in large individuals, and between, ‘one-third to one-quarter in small ones; (@) eyelobe less strongly curved, making an angle of about 160° with the frontal lobe; short, extending for two-fifths cephalic length in small individuals, a little over ‘one-third in large ones (text-ig. 2b); terminates opposite Ip lobe; (©) postocular cephalic length is one-fifth cephalic length (text-fig. 24); (/) interocular area is broader posteriorly; (C2) genal angle slightly more advanced, distal part of posterior border being concave forward (PL. 69, fig. 10); (h) exoskeleton coarsely reticulated (PI. 69, fig. 15); (@ thoracic pleurae (to fulerum) narrower anteriorly, being less than one-third axial width (PL 70, fig. 12). Although having suffered some degree of distortion, the often paraboloid form of the cephalon is considered to be a real, not apparent effect. This is demonstrated by Plate 69, figs. 10-17, the specimen showing no flattening and little lateral compression, burt having a paraboloid form. In Plate 69, fig. 15 the specimen is seen to have been flattened and consequently appears to be a little broader. Hypostome (Pl. 69, fig. 8) is a little longer than broad. Anterior lobe strongly swollen, semicircular; occupies five-sixths hypostomal length. Medial furrow deep, ‘curved gently convex posteriorly. Posterior lobeis very short and crescentic. Posterior border very narrow (sag.) medially, widens laterally into postero-lateral spine. Discussion. Variation in cephalic shape has been noted and may be partly related to local environmental conditions: specimens from the Loch Awe quarry being charac terized by a preponderance of more paraboloid forms, whilst those from Meall a’Ghiubhais tend to be more semicircular. Another variation which occurs between specimens from these two localities is in size; those from Loch Awe tending to be2 PALAFONTOLOGY, VOLUME 21 smaller than the associated ©. Japworthi whilst at Meall a’Ghiubhais the converse is true (Peach 1894, p. 665). These two factors would scem to be interrelated, the cephalon being parabolic in small individuals, becoming more semicircular in the more mature stage (McNamara 1978, text-fig. 4) In large individuals of O. reti~ ‘culatus the genal angle is positioned alittle further forward than in less mature individ uals. The difference between the posterior border in large forms of this species and large forms of O. hamoculus lies in the distal section; in O. reticularus itis concave forward whilst in 0. hamoculus itis convex forward. The shallow epipalpebral furrow (PI. 69, fg. 10 and PI. 70, fig. 1) extends into the frontal lobe as a single furrow and does not appear to bifurcate asin the olenellid Wanneria Bundgreni (Moberg) figured by Bergstrom (1973, figs. 1, 17, 184). ‘The possibility that lapworth? and reticulatus are sexual dimorphs has been ‘examined but because of the above and other factors (see MeNamars 1978) there seem to be no firm grounds for invoking this speculation. ‘One specimen of O. reticwatus has an indented genal angle (Pl. 70, fig. 2). This was not caused by post-eedysial mechanical breakage because the raised cephalic border is intact and follows the line ofthe indentation. Thus the border had been regenerated after the living trilobite had lost its genal spine and postero-lateral genal area, probably from having been attacked by a predator, perhaps shortly after eedysis. Burling (1916) deseribed and figured a specimen of O. robsenensis with healed, broken pleurae in, ‘a position a little posterior to that of the O. reticulatus specimen. The shortness of the eye of O. reticulatus may account for the damage to the genal angle. From the position of the furrow on the distal part ofthe eye lobe which carried the eye surface, it can be inferred that, as the eye surface did not extend to the very posterior tip of the eye lobe, the horizontal range of vision was approximately 80°, ranging from just in front of the genal angle to a point on the border in line (ir. with the middle of the frontal lobe. Thus this species could have been vulnerable to attack from the posterior, so perhaps accounting for the injured parts being in a similar position in both the ‘Scottish specimen and Burling’s specimen. ‘What Peach (1894, p. 665, fig. 2) considered to be the telson of this species is in fact an extended axial spine borne, in O. reticulatus, by the fourteenth thoracic seg- ‘ment. An opisthothorax and true telson have not been observed in this species. ‘0. reiculatus beats very short eye lobes in comparison to many species of Oleneltus, though 0. truemani Walcott, 1913, O. fremonti (Walcott, 1910), and some individuals of 0. gilberti Meck, 1874 also have short eye lobes. Although the eye lobes of 0. truemani are of similar length to those of 0. reticwlarus, they are broader; it also has a shorter preglabellar field. O. fremonti has both short eye lobes and an advanced genal angle, but its frontal lobe abuts against the anterior border. The form from the Eager Formation in Canada referred by Best (1952) to O. gilberti has eye lobes as short as those of 0. reticulatus, a distinct preglabellar field and slightly advanced ‘genal angle. The difference between the two species lies in the extraocular area which is much narrower in 0. reticulatus as the glabella is proportionately broader. Poulsen (1931) referred a specimen of O. reticulatus to Wanneria nathorsti Poulsen (1932), from the Ella Island Formation, East Greenland, but W. nathorst! possesses a much broader border and lacks a preglabellar field, ‘0. gigas is considered to be a large specimen of O. reticulatus. The specimenCOWIE AND McNAMARA: OLENELLUS o (GSE 467) figured by Peach (1894, textsig. 1, p. 667), Walcott (1910, pl. 40, fig. 1), ‘and Lake (1937, pl. 35, fs. 3) has promineni reticulation, short, gently curved eye lobes, the posterior tip of which was probably inline with the Ip asin 0. reticulaus ‘The anterior ofthe specimen is very poorly preserved thus the dimensions ofthe pre- slabellar field cannot be ascertained (Oleneltus hamoculus sp. nov. Derivation of name, Hamus—hooked; culus—eye, Holotype. Compete ephslon, GSE 13302 (1. 70, 8g.) fom the “Puc” Bods, Lower Cambrian, Loc ‘Awe, Sutherland. The locality a roadside quarry 170m E10" 8. of the north-eastern net of Loch Awe C251 158), Dimensions ofhtoype. nm Sagital length of cepalon 09 ‘Maximum with of eephalon 180 Sagittal dimension of prelabellar Held 8 Sagittal length from anterior cephalic margin to anterior gab furrow (3p) $3 Eapital distance rom poster ip of eyelobe o pore border ofeepbaion 29 Poratypes. Six cephala GSM 102276, GSE 13305, GSE 13300, GSE 13298, GSE 13308, GSE 13307, same hoviaan and leality as holotype Matera, horzon, and locality. Twenty-five well-preserved and aumerous fragmentary cephala are known fiom the type locality ia the "Fuso Beds at Loch Awe. Diagnosis. Cephalon generally sub-ellipsoidal; anterior border almost transverse to gently rounded ; enal angle advanced ; distal part of posterior border convex forward. Frontal lobe is well rounded and approximately same width as occipital ring (text- fig. 3a); posterior of frontal lobe less than half cephalic length from anterior border. Eye lobes short, strongly curved, and broad, Description. Cephalic outline varies between sub-semicircular, particularly in young forms (Pl. 70, igs. 4, 6) and sub-elipsoidal in more mature forms (Pl. 70, fig. 3); anterior border nearly straight adaxially; bordered by narrow, rounded rim which flattens and widens only slightly towards genal angle. Posterior border less convex than anterior border; directed almost straight out from axial furrow then, half-way to genal angle in mature specimens (Pl. 70, fg. 3), slightly les than half-way in small cephala (Pl. 70, fig. 6), directed sharply forward in a forwardly convex curve, forming ‘an intergenal angle of approximately 150°, the genal angle thus being in an advanced position level with or generally forward of the occipital furrow. At intergenal angle there is a thickening of the border which, in some specimens, develops into a small posteriorly directed node (PI. 70, fig. 9). Genal spine half as long as cephalon; con- {inues line of postero-lateral border. Glabella ‘hour-glass’ shaped, frontal lobe being of similar width (ir) to occipital ring (text-fig. 3a); slightly vaulted transversely; ‘occupies one-quarter width of cephalon. Posterior of frontal lobe situated at more ‘than half cephalic length (sag.) from posterior of occipital ring. Anterior frontal lobeos PALAEONTOLOGY, VOLUME 21 declines into a short preglabellar field which occupies slightly under one-tenth total cephalic length nlrge individuals, and slighty more dan one-tenth smal ones (PI. 70, fig. 4). Glabella bears three pairs of glabellar furrows: 3p furrow trans- glabellar; directed gently forward abaxially, then strongly recurved back around lateral extremity of 3p lobe; 2p furrow set close to 3p furrow, though itis less strongly recurved distally: Ip furrow, which is only faintly transglabellar, is set further back from 2p furrow than that is from 3p furrow. Flat central glabellar area, one-quarter posterior-glabellar width, is present between gently convex Ip lobes (PI. 70 figs. 5,8). Occipital furrow is similar to Ip furrow, though it is not transglabellar. Occipital ring as long (sag. and exsag.) as Ip lobe; bears a prominent node postero-medially (PL 70, figs. 3, 9). Posterior half of ring lies behind line of posterior border. From, postero-lateral part of frontal lobe, eye lobe arches out strongly, making an angle of ‘About. 140° with frontal lobe; it is short, occupying one-third of cephalic length (text-fig. 26), and broad; it has a strongly curved outer margin but a less strongly curved inner one; posterior tip recurves towards glabella, being situated opposite mid-point of Ip lobe (PL. 70, fig. 9). Post-ocular cephalic length is one-fifth total ‘cephalic length (text-fig. 2a). In one specimen (PI. 70, fig. 9) there isa faint intergenal ridge extending to intergenal angle, Epipalpebral furrow well developed, particularly jn mature specimens; continues faintly into frontal lobe (PI. 70, figs, 6, 9). Inter- ‘ocular area small, slightly raised above extraocular level, which is wide, almost flat, gently downsloping to narrow border furrow. Other parts are unknown, Discussion. O. hamoculus is distinguished from 0. lapworthi in the following ways: (@) the cephalon tends toward a sub-llipsoidal shape, whereas itis sub-parabolic to sub-semicircular in O. lapworthi; (b) the posterior border has a sharp intergenal angle with concomitant well advanced genal angle, whereas it is straight out to much more distally placed intergenal angle in O. lapworthi. The genal angle is hardly advanced in O. lapwrorthi; (©) eye lobe shorter and more strongly curved. EXPLANATION OF PLATE 70 [All specimens from the Fucoid Beds except F1 (Bi. 10) Figs 1, 2,12, Olena retclaes Peach, 1894. 1, GSE $372, internal mould of cephalon from Meal! “Giubhais, Kinlochewe: » 2 Figured by Lake 197, pl 34, Bg. 2. GSE 1330, internal mould of phan roan Loca Awe quarry, nest Fochaadamph; sowing # healed wound of tech WB's. 12, GSE S35, two incomplete dos exosteletons om same lcaliy a fig. 1 2. Peach 194, pl. 30, fg 5 Figs 3-2, Olimar hamoculus 9.90% eral moulds of ephal rom Loch Awe quary, at "Zamph. 5, GSE 13300 ho}ype, 3.4, GSE 1305, paratype; 5. 5, GSE 13M, paratype, <4 {6 GSE 13300, paratype <8. 7, GSE 1307, paratype: "3. 8,CSE 13298, paratype; 228.9, GSM 102285, paraypes 425, Figs, 10,11 Ole intermedia Peach 1894. 1, FU, slice incomplete cephalon from the Salterla ‘Ga Skiag Bridges 7. Tl, OSE $367, internal movld of eephalon rom Meall ¢Chiubhals, Kine leche: boloype» 6. Figured alo by Lake 1957, p35,PLATE 70 ‘COWIE and MCNAMARA, Olene!lus(0. hamoculus is distinguished from O. reticulatus inthe following ways: (a) the eye lobe is more strongly curved (contrast Pl 70, figs. 2, 3); (®) the frontal lobe is more rounded anteriorly and its posterior is situated more anteriorly; (©) the genal angle is in a more advanced position, the distal part of the posterior border being convex forward, not concave forward as in O. reticulatus. A species of Olenellus which is morphologically similar to 0. hamoculusis 0. eager- censis Best, 1952, from the Eager Formation, British Columbia. It compares in having the well-advanced genal angle, short, hooked eye lobe, “hour-plas’ shaped glaella, and almost transverse anterior border. The differences lie in the preglabellar field, Which is not present in 0. eagerenss, and the frontal lobe, which is not as strongly swollen in O. hamoculus. The similarities between these two species is probably 4 reflection of their occupation of similar niches (McNamara 1978) and does not indicate any inherent close genetic relationship as they were geographically widely separated (Cowie 1971, figs. 1,4) Similarity can also be noted beiween 0. hamoculus and O. mohavensis (Crickmay, 1933) from Lower Cambrian shales in the Mohave Desert, California, U.S.A., which hhas slightly longer and less strongly curved eye lobe; shorter preglabeliar field and frontal lobe almost touching anterior rim, even in immature individuals. (Olenettus intermedius Peach, 1894 Pe 7, 10,11 1894, Olenelius inermedias each; pp. 665-668, pl. 32,7. 1910 Olerelis uermeds Peach Walcot, 9.332 1837 Oleelha? intermedius Peach; Lake, pp. 244-245; pl. 35, fg. 1, non 2 “Holotype. Holotype is by monotypy. An almost compe cepalon, GSE $367, fom the ‘Fucoi” Beds, ‘Lower Cambrian Malla'Ghiubhas, Scotland, figured by Peach 1894, pl. 32,7 Dimensions of holoype as Sagita length of cephalon 30 Minimum width of expan 7 Sagittal dimensions of preglabela eld o1s Sagital length fom anterior cephalic repion to anterior glabella furrow (3p) 13. Exsapital ength of ee lobe 05 Exsigtal distance frm posterior tip ofeye lobe to posterior border ofeephalon 1 Material hori, nd localities. nation othe bolype one other eephalon is known from the ame horton and locality as the holeype. Th cepalon which Laks (1937) rleted to this specs 3 stall rmerapid cephaon (1-2 mm, sata! length) of O. rerum. A third, iced, specimen (PI 70, 10) [Skrown fom the Selerefle Gri at Skiag Bridge Conti.) Diagnosis, Anterior margin almost transverse; eye lobes short; preglabellar field very short; frontal lobe short (sag.); extraocular area narrow: intergenal angle close to axial furrow. Glabellar furrows nearly transverse; none are clearly transglabellar, bbut sometimes a very faint connection across the glabella can be seen. Deseription. Cephalon hexagonal shaped, broader than long; border almost trans- verse anteriorly; genal angle advanced, being in line with Ip labe. Intergenal angle(COWIE AND McNAMARA: OLENELLUS on set approximately one-third distance from axial furrow to genal angle; bears a short spine which continues as the raised intergenal ridge which traverses cheek in a gentle ‘curve and meets axial furrow by Ip lobe; posterior furrow which demarcates this intergenal ridge more pronounced than anterior one. Preglabellar field very short, being one-twenticth the cephalic length in the holotype (PI. 70, ig. 11). Glabella widest anteriorly across frontal lobe, narrowing at 2p furrows, then ‘widening slightly to occipital ring. Frontal lobe short, occupying two-fifths glabellar length. 3p furrow curved: 2p furrow nearly straight and transverse; Ip furrow slightly ‘curved, running a little backward adaxially; none of the furrows is transglabella. ‘Occipital furrow transverse and shallow medially, runs slightly forward abaxially and is deeper. Occipital ring bears prominent occipital node. Eye lobe arches strongly from the frontal lobe but is gently curved and short, occupying alittle under one-third cephalic length; posterior tip in line with Ip furrow. Interocular area is slightly swollen. Extraocular area narrow: flat close to eye lobe, gently declines both laterally and posteriorly. Marginal furrow is moderately impressed. Discussion. Walcott (1910, p. 332) considered O. intermedius to be an immature form of O. lapwortht on account of the advanced position of the genal angle. The young forms of 0. lapworthi 2-3 mm in length, though tending to have a more advanced genal angle than the mature form, possess a long eye lobe which, as in more mature forms, reaches to the occipital ring; 50 O. intermedius cannot be considered a young form of O. lapworthi. Lake (1937, pp. 244-245) placed the species questioningly within the genus Olenelfus and included within ita specimen which had earlier been figured by Waleott (1910, pl. 39, fig. 8) and considered by him to be a young form of 0. reticulatus. The specimen, though broken and very poorly preserved, has a longer «eye lobe than 0. intermedius, the posterior tips being in line with the Ip fobe; thus, as Waleott suggested, it probably represents an immature form of O. reticulatus Both Walcott and Lake either questioned the validity of this species or were un- certainastoits true generic position, but the discovery of two further specimens would “appear to validate the species. It is most closely related to O. hamoculus which has a similarly advanced genal angle but differs in possessing shorter, less strongly curved eye lobes, more transverse anterior cephalic border, and shorter (sag.) preglabellar field Bristolia bristolensis (Resser, 1928) and B. insolens (Resser, 1928) from the Lower ‘Cambrian of Nevada similarly have a short eye lobe, glabella impinging against the anterior border, and almost transverse 2p furrow, but the anterior two pairs of labellar furrows are transglabellar, the eye lobe lies much closer to the glabella, and the genal angle is in a far more advanced position. SIGNIFICANCE, PALAEOGEOGRAPHIC AND BIOGEOGRAPHIC RELEVANCE OF THE FAUNA ‘The olencllid fauna of the Fucoid Beds is undoubtedly a North American Pacific Province association and the zonal significance is probably to date the ‘Fucoid Beds’ ‘as in the younger part of the Lower Cambrian—the Boriia-Olenellus Zone (Fritz 1972). Deposition of the Fucoid Beds will have thus been in the north-western shelf sea (lextfig.on PALAEONTOLOGY, VOLUME 21 WORTH ATLANTIC [SRS Seine REGION sD > US Part ~ palinspastic J \ Inap for the Pati FTS Piaovince CAMBRIAN x ae Mint * of \ SN ; # ‘iB Yeo! PROVINCE] ‘xr, 4 pata tap forthe Cambrian Period aa — a region. The brsier tetwoun the Pacibe nd Arado-Balts Provinces of the Oleelhd Realm, posibly deeper water ofthe Tapets Oceat sof unknown Width abd not intended tbe to any scale. ‘The faunal provinces of the Lower Cambrian of arctic North America, Greenland, Spitsbergen, and Scotland have heen discussed by Cowie (1960, 1971) and Palmer (1569, 1572, 1973, 1974), The Olenelid Realm is divided into: (i) the Aeado-Balte Province with characteristic trilobites which inelude Callavia, Holmia, Kjerulfia, Strenuaeva, and Strenuella, and (2) the Pacific Province with characteristic trilobites which include Bathynomus, Bonnia, Bonniella, Nevadia, Nevadella, Olenelius, and Protypus. Tn text-fig. 4 a palacogcographical reconstruction has been made for the Cambrian Period which may be close to the Lower Cambrian situation. On this interpretation the Acado-Baltic Province was separated from the Pacific Province by the ancient ‘ocean of Tapetus (Harland 1973; Harland and Gayer 1972). The faunal provinces ‘Would then lie on either side of this geographical barrier which prevented mixing of ‘elements of the two provinces. North-west Scotland would have been on the north- ‘west side of Iapetus with Greenland, Spitsbergen, and maritime Canada.COWIE AND McNAMARA: OLENELLUS a Acknowledgements. The athors thank the following for assistance at various stages ofthe work: Miss ‘ACP. Gon for valuable asance in Uibrary laboratory, and feld investigations; De. R. h. Wison, Mi-P-J Brand (G'S Esinburph), and Dr. A. W-A.Resbion (GS. London) for museum tu fits, ranging loans, end general encouragement: Me. R, Godwia, Mrs A. Grepory, Mrs. J. Bess, and Mrs |B Hilor peparingthe photographs, the lusrations, andryping the manuserpt; Profesor HB Whiting ton kindly ead the manvscript nd encouraged the reversing te use of fais by KJ. MeN, abd Profesor D. L. Dineley for provision of apace, equipment and asstance at Bristol. Thanks are given to {he Natural Environment Research Counl fra research pant for W.C) REFERENCES ‘enasrnont, 2 1973, Organization, if, and systems of tloites. Fale and Stra, 2, © pp, Spl ‘Oso. es, xv. 1982. Two new species of Ole rom British Columbia, Trans. R. Soe. Can. 46,2. 3,13-22, 1p rmcitvos 1861, Palacoroe fst Vo. 1. Geol. Sur. Can 1-24 ‘WIE SH U.,DAWSON,) GALLAGHER J, OSTLE,D. LAMBERT, Rand LAWSON x. 196. Potassium "oh sediments the Cambrian of Norawest Scotland. Tran nun. Meal (Sot. BY” Appl. Barth e175, B10, saxo, P1965, Now Lower Cambrian fos localities in NW. Ssotland. Set. J. 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Revised typescript recived 18 June 1977 ‘Australia