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EXOSKELETON, MOULT STAGE, APPENDAGE MORPHOLOGY, AND HABITS OF THE MIDDLE CAMBRIAN TRILOBITE OLENOIDES SERRATUS by H. B. WHITTINGTON [AWHRACK. The external surface and morphology ofthe exoskeleton of Olenoides serratus is describe the ‘xoskslta layer hasbeen replaced by ite and clone. The pleura curves down distally, there 0 fulral process or socket, but a Mange assis in articulation. An early moult tage has the exoskeleton unminerlized {nd wrinkled, and was dexrbod as Nathorstiatranstons. The type i redscribed: «second specimen is ofthe fxoskeleton only. A new rconstraction othe biramous appendages given. TRe outer branch was attached {o the coxa so that it was extended close below the ventral cuticle ofthe body the function was probably fespiatory The inet and ventral surfaces ofthe cox, and the venta side of the outer branch Were spinose. "Tes limbs were adapted Lo size sot prey, squeeze, and shred it and past forward to the mouth; a manner in which these activites could have been effected oulined. The peobable musculature of the hmbs and a ponsble gat are deveribed: this ait unlike other modes of progression in tate that have Deen ro Dosed, How 0. serratus may have jumped off the bottom and drifted is suggested, the swimming powers were probably feeb i could have plowghed shallowly in sot sediment BY alterate flexure and extension of airs [St Tinbs i could have dug in search of prey oF for concealment; a Rusophyeus type of truce may hav ‘sulted trom sochsetiviyO.seratn i ths considered to have Ben a benthic peda and Scavenger. NO Plausile lit movements by which it could have mae a Crane trace are evident: how ths take WIS imide isan unsolved problem ‘Tue earlier description of appendage-bearing spesimens of Olenoides serratus (Whittington 1975) included a new reconstruction of the nimal and a brie? discussion of habits. Here additional details ‘of exoskeletal composition and morphology are given, including those of unusual articulating devices in the thorax. The rare early moult stage (Nathorstiatranstans’) is also redescribed. Further pre ‘aration, new photographs, and drawings have enabled a more detailed restoration ofthe biramous Appendage. The limbs of . serrarus are now better known than those of any other species, and the basic limb movements here portrayed are those regarded as plausible by Manton (1977, pp. 39-53). Certain of these activities produce characteristic trackways. Studies of trace fossils tributed to trilobites (Bergstrém 1973; Birkenmajer and Bruton 1971; Campbell 1975; Crimes 1970, 1975; Martinsson 1965; Osgood 1970, 1975; Seilacher 1955, 1959, 1962, 1970) are largely ‘concerned not with such relatively simple tracks but other and more complicated ones. These trace fossils have had to be interpreted in the absence of any incontrovertible evidence as to what animal ‘made them, but suggestions have been derived from them of the behaviour and limb movements of trilobites. Certain ofthese suggestions are examined here with reference to O.serrarus, and against the background of Manton's work, Ia O. serrane it appears that the filaments ofthe outer branch of the appendage cannot have been used in excavating or brushing away sediment, i. cannot have produced the fine scratches attributed to such an activity. Much attention has been devoted to the {race fossils Rusophycus and Cruziana. 1 appears plausible (text-fg. 14) that 0. serrarus could have smd the former type of trace. On the other hand, no plausible solution has been advanced for the problem of by what imb movements was the trace fossil Cruziana made. I cannot devise one, and {an see no plausible activity of O. serratus that would have resulted in this type of trace. No attempt is made here to re-assess any trace fossil. The suggestions made do, however, call into question interpretations of trace fossils that have gained some acceptance; greater caution is needed in such (Pty, Ya Pat, 190 ph. 11-204 17-285 m PALAEONTOLOGY, VOLUME 23 interpretations. 0 serratus belongs to a relatively small Middle Cambrian group that had few thoracic segments and the cephalon and pygidium are of similar size. Caution must likewise be used in drawing generalizations ftom this single species. LOCALITY AND STRATIGRAPHICAL HORIZON Specimens referred to herein came from C. D, Waleot’Pasllopod hein the quay that he opened in 1910 {or collecting from the Burgess Shale. This quay i stused on the ridge between Wapta Mountain and ‘Mount Field, at an elevation of approximately 7286 m (7800 M48 km (3 mils) north of Fi, southern British Columbia, Canada. Frit(1971) described the geological seingof the Shale and showed that he quarry vas thin the Pagetia Bootes Faunule ofthe BatvuracusBrarina Zone of the erly Mile Cambrian, The Prhsllopod tod is 23 m (71.7 in) thick, but the exact boron from which Walsots specimens came fe 10% known Parties led by the Geological Survey of Canada collocted from the quarry in 1966 and 1967, and ‘measured the levels within the Phyllopod bed ftom which thei specimens were collected (Whitington 1971), Tn the explanations ofthe pats bere, specimens collected by Walcott, now in the U.S. National Museum, are recorded as from the Payllopod bed. The level vith this bed from which the Geoloexal Survey of ‘Canada's specimens came i given Deals of other specimens and Unie oocurtence ae in Whitington 1978, pp. Ut-103, fg 1 METHODS AND SYMBOLS Photographs. Those showing the exoskeleton (I 17, fg 2; Pl. 18 fis. 1-3) have been taken wing normal ‘orthewes lighting (orth at the top ofthe pat), the specimen lghly coated with ammonium chloride sub Iimate- All other photographs were mado om panchromatic lm in wlravolt radian, after focusing is ‘ordinary light. In Some (PL 19, figs 3 4; PL 20, fas. 1,3; PL 22, i 1) he radiation was dieced at about 50° to the horizontal, the direction varied to bring out particular features. The diection is given in each explanation. In the romainder the radiation was directed at about 6510 the horizontal, andthe specimen Uted to about 12°; this gives maximum reflection (fom the surface ofthe specimen. In all except Plate 22, fig. 3 (referred to as ‘refed, the spocimen was covered With a thin fm of dsied water beneath & glass slip. The effect is fo enbance defnion and contrast between matrix and specimens these photograph re fefered wo sy ‘under water The part of specimen is taken (o te that which shows the animal i dors spec, photographs of counterpart only ate s0 specified n the explanations. Testsigures. Two of the ex-igures which explain my interpretation of the specimen are refered to as com: posite because they combine none drawing, features shown by part and/or counterpart. Symbols se tthe fextligues areas follows: {A axis of promotor-remotor sing of cota abr, articulating halsing: am, anterior margin; an, antenna. ap, appendage: ax, ata rng; ch, coxa- body Juneion ce, cores; cox, coxa’ dy dapressoe muscle’ dl it eof gil branch: do, dublure;es, dor excskeleton flexor muses; fanges fo, ol: sul indiaing ail (outer) branch: en, snathobase: gs, genal spine: G-S.C.- Geological Survey of Canada’ hinge joint infling: im, inner margin: pt interpearal furrow: i inner surface: L, as prefix denoting let side of imal; ovat muscle; lateral ps; m, mould of exoskeleton ma, margin 0, outer surface p, pot Joint: pl, pleura: pf pleural furow: pls, pleural spine; py pysicum, sopnents numbered 1py, 293, te ys borer spine of pygiium: pul proximal lobe of outer branch: Ras prefix denoting ight ste of animal { thoray, sven segments numbered It, 2, ete: USIN.M. United States National Museum (now National ‘Museum of Natural Histor), venta; ve, Noi i rock: % aus of articulation in thoras. TIS, Arabic numerals used 1 denoe successive horace segments axl rings of pyidium, plural arrows, podomeres of leg branch, ste Also ase to denote series of biramous appendages beginning with the most Enterior shown by the spasmen, when it was ot the most anterior in the animal TTXV, Roman numerals used fo denote sucesive bramous appendages na seis, begining With the most anterior in elation to the animal, no the specimen chars represent minal searps in the specimens, produced when the rock was originally split, oF by preparation. The ine runs slong the upper ede ofthe vearp, the hachures dieced dow slope fom thine "The scarps un along the broken edges of fagmenls of the exoskeletal ayer, and atthe changes in level which separate pars ofthe body. Inthe drawings they indicate the rlatve lve of ove pat ofthe body to another. ‘Suppl indicates the position of rides and furrows, other uss Of sippe are explained india. WHITTINGTON: OLENOIDES SERRATUS m EXOSKELETON ‘The outer surface of the exoskeletal layer preserves in detail the original form (PI. 17, fig. 1; PL 18, figs. 1-4). Granulation is coarsest on the posterior half ofthe axial rings, on the glabelia and cheeks, and on the pleural ribs of thorax and pyaidium:; furrows are finely granulate to smooth. At the ‘cephalic borders, and borders and spines of thorax and pygidium, the granulation grades into raised cusps which distally merge to form raised lines (PI. 18, figs. 1. 4) On the anterior slope of the Slabella and hypostome similar lines are formed (Whittington 1975, ig. 18, pl 1S, igs. 1-3). The Step searp edges ofthe cusps face inwards, and so are opposed along the edges of the spines. On the Ventral side ofthe spines and along the pygidial doublure are raised, anastomosing lines sub parallel to the margin, The inner Surface ofthe exoskeleton was smooth, as shown by’ the mould ‘where the exoskeletal layer has been stripped off. The broken edgs of the exoskeletal layer (visible for example where it descends vertically into the oublure, Pl 18, ig. 2, ex) appears black and shiny. A thin section Was made across the posterior pysidial border of GS.C. 61375, Waleott Quarry, level 6 18 75 in, to 7 ft 3 in. an incomplete specimen showing portions of appendages. The exoskeletal layer is composed of minute flakes of ‘mineral, the flakes approximately at right angles tothe surface. Electron microprobe analysis ofthe layer showed the presence of two distinct phases, an outer ofilte (pet cent by weight, MgO 485, ALO, 31:32, SiO, 37-20, § 0418, K,O 5-19, CaO 015, FeO 1393, total 9282, minus water), and an inner of chlorite (per cent by Weight, MgO 993, ALLO, 2468, SiO, 2551, $ O48, KO 067, CaO, 0:15, FeO 27:56, total 87-98, minus’ water). A fragment of the exoskeletal layer removed from GS.C. 61374 (PL 18, ig 3), gave the X-ray powder diffraction pattern of illite I is assumed that the exoskeleton was originally mineralized with calcium carbonate, then replacement by ile group ‘minerals occurred at some stage in diagenesis, perhaps at an early stage in an anoxic environment ‘Analyses quoted by Conway Morris (1977, p. 5) indicate thatthe soft parts of fossil worm fom the Burgess Shale are preserved in a film composed of calcium aluminosilicates, formed at some stage in diagenesis. Piper (1972, p. 173) remarked on the presence of diagenetic chlorite in portions of the Burgess Shale. ARTICULATION OF THE THORAX ‘The earlier description of G.S.C. 34694 (Whittington 1975, pp. 119-120, pl. 20, figs. 1, 2.4 pl. 2h, figs. 1, 2: textfig. 22) was primarily eoncermed with the appendages. Further preparation has revealed more completely the cheek (Pl. 18, fg. 4), and also the structure of the thoracic segments, including thearticulating flange briefly alluded to earlier (Whittington 1975, p. 129). The lange fitted below the posterior pleural band of the segment in front, that of the frst thoracic segment below the posterior border of the cephalon. Where the outer surface of the dorsal exoskeleton, or the ‘mould of the inner surface, is not broken, the flange is concealed (eg right side of pleurac 5 to 7, I-18, fg. 3; Whitsington 1975, pl. 1, ig. 2; pl. 9, fig 1). In 34694 the outer surface ofthe flange is exposed by the split or by preparation (PL. 17, Bg. 1; textsfg. 1; PL 18, fig. 2; text-fi. 2). The mould ofthe inner surface ofthe exoskeleton ofthe pleura, where exposed, lies only slightly above the outer surface of the exoskeleton of the succeeding fang, so thatthe two surfaces may in places appear to pass one into the other: in other places the sight difference in level i lear. The posterior marin of each Mange is delineated by a narrow groove, and the flange is granulate. The flanges of the pygidium and seventh thoracic segment have been completely exposed by excavation (PL. 18, fig 2), in more anterior segments the ange is partially covered by the posterior pleural band. The abaxial margin is directed inwards and forwards, making a slight angle with the outer edge ofthe pleural spine. The flange is widest (exs.) abaxally, slightly narrower over most of its extent, Proximal narrowing progressively as it curves down into the axial furrow. In the base of this Furrow it merges into the outermost portion ofthe articulating halving. The ventral exoskeleton of the pleural spine extends inwards to form a band along the abaxial edge of the pleura, which narrows forwards and dies out, and does not extend under the flange. Postero-lterally the inner