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PAEDOMORPHOSIS IN SCOTTISH OLENELLID TRILOBITES (EARLY CAMBRIAN) by KENNETH J. MCNAMARA Ansrasct: Ole (Oaies) armates Peach, 1894 hs en ret and interrted a ving evened by tedomorphoss By compares vin lcd catogeis, 0. (Oli) hams (lets) marmot an 1B (Ott) reelanr a rom the ‘Faso Beds ar ewe comer to have eae by pasomorphoss ‘om (Onl pert Prstomorpbesimaybavccurednesponsetosdapsiontssonere igh onee Stet nals acre edna mean was peogenes wich may have Bem ged Ye ‘tah lmportures of the shalome mtr enoonent wich th lll ao cpl DURING a restudy of the curious litle olenellid trilobite Oleneltus (Olenelloides) ‘armatus Peach, 1854 from the ‘Fucoid’ Beds of north-west Scotland, it soon became ‘apparent that this species, far from being genetically very different from the other Four species of Olenellus which are also present in these beds (O. (Oleneilus)inter- ‘medius Peach, 1894; O. (Olenellus) reticulatus Peach, 1894; 0. (Olenellus) apworthi Peach and Horne, 1892; and 0. (Olenellus) hamoculus Cowie and McNamara, 1978), is closely related to them. The development of diverse olenellid species is not dis- tinctive of the “Fucoid” Beds but isa feature of many other olenellid-bearing Lower ‘Cambrian rocks. This profusion of forms has caused many problems in olenellid taxonomy with new genera and species being erected and placed in synonymy with sgreat frequency. It is hoped that this study will not only help to elucidate the evolu- ionary mechanisms operating within the Scottish olenellids, but also assist in explaining some of the vagaries of olenellid taxonomy in general In his study of O. (Olenelloides) armatus, Peach (189, pl. 32, figs. 1-6) illustrated only four specimens. He used one of these, the part and counterpart of an incomplete, articulated individual, asthe basis for a reconstruction which has become the standard representation of the species (Raw 1957; Moberg 1899; Poulsen in Moore 1959). As the specimen on which Peach based his reconstruction is very poorly preserved and is distorted, Peach's drawing is misleading in its representation of the number of thoracic segments, orientation of the spines, and other minor features. A new recon- struction based on all four known articulated specimens is presented (text-ig. 1). (0. (Olenelloides) armatus has been regarded by many previous workers (see below) as either ‘larval’, ‘immature’, ‘primitive’, ‘degenerate’, or ‘aberrant’. Many even doubted its specific validity, whilst Hupé (1953) considered it sufficiently distinct ‘to warrant placing within a monotypie subfamily. In the present study itis hoped to clarify its taxonomic situation. Terminology. This largely follows Harrington ( Moors 195) except that proeranidial spines preferred to'pertnigenal spine" anditergenal spine fmetafngeal spine’ The tes inergenal ge sintrodueed to deeribe the pleural extension athe preociptal pment Palmer (987.10) flowed ia usiog the {Panos Yl 2, a3, 18 6-65, 7 a6 PALAEONTOLOGY, VOLUME 21 term ‘interoelar area’ forthe iene" of Hupé 19822, p. 117) The genal aren outside the eye lobe is termed the “xtaoeaar area" Th je lobe separated inn the aleror and posterior ee edge, between Which isthe ‘epipapebal furrow’ Hypostome terminology follows Palme (1957, p. 107) “Pacdomorphoss' was defined by de Beer (1958) as ‘phylogenetic effets produced by introducing youthful characters into the line of adults’, whist ‘proenesis a term introduced by Gard (1887) t0 ‘Sseribe precocious sexual maturation ofthe repextutive organs ube the organist sil in the Soni ‘ofa lava. De Beer (19S, p. 63) included progeness with nooteny, asthe end reul eention of lava Features nthe adult sage, the same. He alo nstudad pacdogencis (von Bas 1855) with noon, bul the mechanisms of both progeneis and paedogencss are very diferent from that of note resulng in variation inthe maximo sie ofthe adult. In neoteny the preadult ate of development ofthe organism {Gncluding its sexual development is delayed or arrested, resulting in retention of tral characters i he ‘adult Doe toa peolonged period in the pre-adlt stage, te pre-adul rate of growths prolonged, con ‘Scqueatly the neatenousadult sarge thanthenon-seoteous adult Pacdogenes rested. Oren which reproduce ina val stage, bt parthenogenetic asin some Coleoptera (2 development continues t normal pace until precocious ‘ered inthe pre-2dult stage only allows a small degre offs jen growth, wih consequence hat he “ult peogeneti form cannot reach sarge sie asthe nonrogencie form. Wheres pssously este tas been thought tobe the only machanim producing puedomorpic evolution (de eer 1988, p68) vidnce i prsuced hereto show that progentts as becn the poedomorphic mechanism inthe oleae Telobite in north-west Seotnd” Gould (1977) has recently, dependent, come (o smi contusions ‘concerning the pacdomorphic processes SYSTEMATIC PALAEONTOLOGY Family oLeNELLIDAE Vosdes, 1893 ‘Subfamily OLENELLINAE Vogdes, 1893 Tinel. oLeNeLLomivat: Hupé, 19536] (Genus OLENELLUS Billings, 1861 ‘Subgenus OLENELLOWES Peach, 1894 ‘Type species. Olen (Olenellodes)armarus Peach, 1894; fom the -Fucold” Beds (Lower Cambritn, ‘Meal s'Ghisbais, Ross and Cromarty. Scotland, ‘Remarks. Although Peach (1894) erected Olenelioidesas.a subgenus, he did not present the species as 0. (Olenelioides) armavus’, bt as ‘Olenelloides armatus’. Consequently, later workers used the name in a generic sense without justification. It will be shown that although 0. (Olenelloides) armatus is morphologically very distinct from con- temporaneous olenellids from the same horizon, genetically it was probably very closoly related to them, Thus Olenelloide is retained inits original sense asa subgenus of Oleneitus. Olenellus (Olenelloides) armatus Peach, 1894 Phe Th fs. 1-9 tex 1194, Otenetis(Olentowes)armanus Peach: pp, 648-670, pl. 3, Hes. 1-6 1897 Olenlodesamars Peach; Becher, p91 {9 i910 192). Olenelodes amatus Peach Raymond, p. 12. 1925. Olnelliesermatus Peach: Ra, p23 1937 Oineble amar Pah: Lake. 286 01.38..79. 1937 Olnelloidesarmatus Peach: Raw, Bp. $2 S83, 30. MeNAMARA: PAEDOMORPHOSIS IN OLENELLIDS on 1938 Olnetoidesarmatus Peach Lake, p. 249-250 pl. 36, fi. 1 1842 Olonetloiderarmatus Peach: Storer, pp 65, 105. 19530 Olonelloies eran Peach; Hupp. 125, 19536 Olonetionesormatus Peach: Hope. p75. 1957. Olenellodesarmatus Peach; Palmer, pp 12-122 1987. Olenetodes amas Peach! Raw, pp. 149, 130, 158; text | 1971 Olontoidesarmatus Beach: He, p63, 71,79. 1973 Oleneledes armas Peach; Bergtcen, p33 Lectotype Hecsn designate; an almost compete cephalon (GSE 472) fom the ‘Pacoid’ Beds, Lower Cambrian, onthe nomthers slopes of Mall a Ghiubs, Ross and Cromarty (locality 6 in wextfg. of ‘Cowie and McNamara 1973); red by each (1894, pl 32, fg. 4) and Lake (193, pl. 36, fg 1); figured Plate 7, fig, heen ‘Materia, loaiy, and horizon, The only materia avilable iin the Geological Survey Collection made by Mr. A. Macconochi inthe later part ofthe nineteenth cnt. Receat tempts have ald to reveal the bsewhich yielded 0. (Olenelloie) rman, Thespecisens are maily housed isthe lnsitutof Geologic Science in Edinburgh (GSE), a small number also being housed inthe Institut of Geological Sciences in London (GSM). Four articulated specimens, thirty-three largely complete and fourteen fragmentary “xphals and to hypostome are own Irom the same horizon apd locality asthe ecotype. The specimens ‘sre preserved as ochreous internal and exeral moulds ns dark, yellow-weatheing shale Diagnosis. Exoskeleton long and narrow; small. Cephalon hexagonal, bearing long, ‘equidimensional procranidial, genal and intergenal spines; genal angle in line wi 2p glabellar lobe. Frontal lobe short, abutting against anterior border. Eye lobe very short and strongly curved. 2p glabellar furrow transglabellar; all glabellar furrows except 3p, directed almost transversely. Interocular area strongly lobate and similar in width to very narrow extraocular area. Hypostome denticulate; posterior lobe large, occupying one-third area occupied by anterior lobe. Thorax composed of nine segments; pleural regions ery narrow (ir. Remarks. 0. (Olenellodes) armatus was adequately desribed. by Peach (1594, pp. 669-670), Walcott (1910, pp. 347- 350), and Lake (1938, pp. 248-249). However, « numberof further features of specific importance can beaded to their descriptions. Theanteriorcephalic border is almost transverse, but arched slightly forward where frontal lobe abuts border (PL. 71, fig. 1). Antero-lateral margin slightly concave and three-quarters as iong a8 anterior margin. Postero-ateral margin strongly concave and equal in length to anterolateral margin. Posterior margin transverse: width (ir), and width of occipital ring, equal to that of anterior xrnc |, New recantation of 0. (Okt margin. 3p and 2p labellar furrows rans Fas, 1