Bollenino delk Societh Pahontohgica Italiana 43(3),2004 rssN0375-7633 383-40r 3 pls. Modena, Dicembre 2004

Cambrian microfauna and palaeoecologyof the Campo Pisano Formation at Gutturu Pala (Iglesiente, SlUf , Itdy)

Olaf EI-Icru Gian Luigi Ptllom TU BereakademieFreibere Universit)r deeli studi di Casliari Geologiscf,esInstitut, Freibirg Diparti-ento"Scienze della Terra

I(EY lrfORDS - Microfossils,Palaeoecologl, Campo Pisano Formation, , Sardir

ABSTMCT - The microfossilcontent of nodukr limestonesof the late Early to Middle Gutturu Pala sectionofsouth*ritrrn Sardinia is describedand discusid with ,ttpttito pakeoeco tigations, the uerticaliuccasion of the sedimentaryeuolution andfossil distibuiion is intqrpretet to at hast deep-subt;dil con, tfaot conditionsa! tle bgginyini of yhe Campo.ri:f"y Formatioln tics, an interim shalhwing during that processu likely. A dramatic facies,toTntt*eit wit6 the onsetof tbe Cam\o PisanoFormation indicat€slt strt by tectonicinstahitity oftht pktfor*, Accompaniel.by modeiaterektiue sea-leuelrise and pakeol frott of the time, thi pirsistintiof onb onepaktott6hgical assembkgeindicates a more5r hssii for' mo,stof the,CarnpoPisano time. Aming the taxa representedin the highly fossiliferoussediments, , echinoderms,b elementsarZ chancelhrilids,hyolithids, ptk7leltih, hiolithelminthids, and bradoriid . is pubtished herefor the fii, tirye. The foaqwing bradoriid and trih_Qitesryciel are newly t Condylopyge aniiqua n.-sp.,and - dzscribedin optn nomenclature- Clavigellusi n.sP.

RASSUNTO - [Microfauna cambrianae paleoecologiadella Formazionedi Campo P serrenrrionale,Sardeena)l - I mimofossili,ontiruti nei ilcari nodukri d,el Cambriaio ini CampoPisano drlk sizio'ie di Gumriu Pala, Sardegnasud-occidentale, sono desritti e discussiT base-didenasliate analisi di facia, dell'eaoluzione-sedimentariae delh distibuzione dei resti I in termini fr uariazioni ambientali che indicano condizioni sub-tidali pocoprofonde alk baie dizioni sub-tidali profonde(probabilmente anche mobo profonde)alk immith"delk stessaforn no inoltre k presenzid; un'episodiointermedio conekbili con un sensibilzabbassamento dzl h La dmiica sostituzione'difacia, che segnal'inizio delh Formazione di Campo Pisano ambienti. Cib b spiegabileinnanLitt tto con lTinstabilitt tettonicafulk7iattaforrna, ilk quale . liaelh marino e'uaiiazioni climatiche indotte dall'assenopaleogeografco +i p_ersistenzn,dura associazionepaleoycghgicg, indica chek deposiT|gnedelk mag'ior Parte delk Formazione di C, no aperto,' sib-tidale profondo,ph o menostabile. FrA i gruppi di org;nismi ,apprtttntati nei sedimentiriccarnente fossihferi predominano tr, inolne prtTt"ii ,o*poirnti minoii qu.ali chancelloridi, hyolithidi, pikg;til;dl,'hy.olithelmintid Formaiione di Cnkpo Pisano e piOOt;cataper k primla uolta ii quZstoarticih. Sono state Hipponicarion ichdusum n. tp.'t Condyloiyg. airdqua n. sp., appartenentirispeniuamente mentreunt nuoaAspecie di n;hbiti coryiexichTdi,ott ibuita i Ctiiigellusi ,. tp., i stata laci

INTRODUCTTON tic shelf environment (Nebida Group) to an isolated carbonate platform ( Group) followed by In southwesternSardinia (Texr-fig. 1) autochtho- the drownitg and cover of the platform by siliciclas- nous Cambro-Ordoviciansediments crop out in the tic sediments (IglesiasGroup; Texvfig. 2). Iglesienteand area (summary monograph: The Nebida Group (Early Cambrian) consists of Bechstadt& Boni, 1994).The Cambriansuccessiop a basal siliciclastic portion with local calcimicrobial- within this suiteis under investigationsince the l9m archaeoryathan buildups (Matoppa Formation) and century.Vhereas the earlywork was mainly focused an upper alternation of shallow marine siliciclastic on palaeontologicaltopics (e.g. Bornemann, 1886, deposits and carbonates(Punta Manna Formation). 1891; Meneghini, 1888), researchchanged later to The overlyi.g Gonnesa Group (Early Cambrian) is sedimentological,facies and other interests. completely composed of carbonates: whereas the The entire Cambrian successionhas a thicknessof lower part (Santa Barbara Formation) is mainly rep- about2000 m and indicatesan evolutionfrom a clas- resented by dolostones, the higher part consists of 384 O. ELICI(I, G.L, PILLOIA

detailed study. Investigationson this unit were focusedin rather subordinateaspects of work on the whole Cambrian successionor on specid palaeonto- logical remains.Followitg that work, the lower unit (Campo Pisano Formadon) representsa condensed carbonatesuccession consisting of more or lessnodu- lar silry limestones(Loi et al., 1995). The origin of the nodularity wasinterpreted by Gandin (1979) and Cocozza&, Gandin (1990) asdue to compactionand early dissoludonprocesses on a tectonic instableand -l+Ft'\fY rr + morphological differentiated shelf, These authors l/e€:+,*,*_t compare this nodular facies to the GibagrabenNiri.**** "Ammonitico Rosso".In contrast,Elicki (2001) Pre- sentedarguments for a depositionin a morphologi- cally rather low diversified subtidal shelf environ- ment. After that author, the nodular texture of the Campo PisanoFormation is a result of complex late diageneticprocesses. Palaeontologicdly, the Campo Pisano Formation was never subjectedto intensive investigationuntil recentyears. Sporadic works came from Cherchi 6( Schroeder(1984, foraminiferaand problematica),from Mostler (1985, poriferidt) and t] cenozoic O Gutturu Pala hom Pillola (1991, trilobites).Since the middle of H #l Mesozoic @ the 90's, the microfauna of the Campo Pisano is investigation by the Freiberg f allochthonousPalaeozoic O Iglesias Formation under University working group. Hitherto, some work on E granites "small m early to Variscan the so-cdled shellyfossils" is alreadypublished : IglesiasGroup 13] ?Precambrian (2002), Elicki et al. (2003) and Elicki 6c 'Wotteby Elicki I GonnesaGroup \l thrusts (2003). A number of further publications %' from there are in progress. Nebida Group faults N Regarditg the higher part of the IglesiasGroup rex'-ngI - 'ffif (CabitzaFormation), some newer work is on the bios- ff:J'E::*#ff,::',::iffi ilti:'i,S"iiii: tratigraphy and sedimentolog;r of this siliciclasdc workingsection at GutturuPala locdiry. unit. Loi et Al. (1995) wereable to subdividethis por- tion by trilobites and graptolitesinto six different biostratigraphiclevels reaching from the early Mid- (Caesaraugustian limestones (San Giovanni Formation). The youngest Cambrian to the Early Ordovician unit of the Sardinian Cambrian is the Iglesias Group to Themadocian).The'depositional envirot -Jttt fol- (uppermost Early Cambrian to Early Ordovician), lowing the Campo Pisanocarbonates is interpretedby rt bi"i"g a carbbnate basal part (Campo Pisano different authors as e deeperbasin setting basedon (Gandin Formati-ot ) and a siliciclastic- upper part (Cabitza sedimentologicaland taphonomiccharacters Formation) that continues into the Early Ordovician et al., 1987; Cocozza6c Gandin, 1990; Loi et dl., Mid-Cambrian to the (T.{-fig.z).. . . 1995). For the late During the last decades,there was an intensive Themadocianhigher CabitzaFormation, in contrast,a sedimerrtiogical and palaeontologicalwork focused relativelyshallow environment was interpretedby Loi on the Early Cambrian units, e.g. the archaeoryaths et al. (1995) becauseof rypicd sedimentaryfeatures. were investigatedby Debrenne, Perej6n, Moreno- The aim of this paper is to presentfor the first Eiris and .Jl."g,r.r, the trilobites by Rasetti and time the microfauna of an entire section of the Pillola, and thi sedimentary facies mainly by Campo Pisano Formation, which was attained by Cocozze, Gandin, and by the working group of chemicalpreparation and by thin sections,and to dis- Bechstiidtand Boni. A summaryof the palaeontolog cusstheir palaeoecologicalrelevance. icd, sedimentary and facies investigationson the The palaeontologicalspecimens described and fig- Sardinian Cambrian, as well as an extensivelist of ured heiein are housedat the GeologicalInstitute of publications was published by Pillola (1991) and FreibergUniversiry (archive number 536),despite the Bechstldt& Boni (1994). material, which is at Dipartimento di The uppermost Early Cambrian to Earty Scienze della Terra ( University; acronym Ordovician IglesiasGroup has receivedonly little DSTCP 23). UMBRUN MICROFAUNAAND PAAEOECOLOGYOF THE CAAIPOPISANO.,. 385

carbonate facies types: m - mudstone, GI Tw - wackestone, f - floatstone q o Ft m T r-.'r' Esc) o tr -=r- €t Fl ka Er -{---f- I clt !iE +, (); ----r=-r _q cl N ..*l - II F u, La o ! (t)c) a Hq tlt U E F-{ F. cl !€ ?a o0 w gs (D (D a iOI DlrDl(D l( o (D (D (D (DlO q) C.P. (DI(D ID .D (-l(D O I-I-I- Fl (D I(D:(Di(D (D (D .D .D olrLe lsirl (D ID (D (D a) l(D I (D -T-^I ft-rur- (D .Dl(t ala It .D (D (D (D .D ol- rD (D (D (D (D )l(Dl(D (D E ID (D (D .D (D .D tr E l- d tr tr GI I A (D (Dl(D (D (D o (D (D (D (D (D o # w l(D m f-. E (, rt-l-T- (D (DI(D (D (D 5 =\J iv O5 a \-=l=r--r--f-- --2'-1-J*,-< -r'--L--/ SF +i:l# qT -<' * \>,-/-\ oo tttl T F. rn = c.t (D .D .D (D iloToJ5 o tr -7' t=z__j_____/ .D ;f-l--t- f. zrt T& (D (D v ol-lolo ct (D I 1l ,l-T-lo_L G' rDl(D - ID ? IO I-IO ! w rl (D h ct tr o GI tr a 'tr d -o tr rt cl rt E (D (D rD crl li E rtl-T't'r'i- OIO Fossil content: cl fr{ (D (Dl{D (D .D U C(t tr (D (D (D (D -b tr m D - - hvolithids li G' (D (D (D O (l lo H .D OIO (D (D (D chancelloriids GI (D (Dl(D (D (D ffi tr Ltallrltal .D l.D oirDlrD .D (D (D brachioPods trr -]1- _l_ -l_ w (D rD Ioiolo tt (D F!i!i!i!il (D I.D (D .D ID (D (D A (D .Di(DlOlO (D (D I tritobites -l(D iD ID .D .D (D I tE f 6E ,tl l-l echinoderms clT E €o E{ w poriferids GI Effi t 58 g lt++f+t+t+t l o A tr o R3 e a GI z GI dolostonedolostor ttu= c) - tl -_F-rE nodularnooularr r limestone t samplingintervals o ;?fil I limestonelimestor L faunalintervals = { ffi sandstonesandstor Ee I claystone/siltstoneclaystor

Text-fig.2 - Geological column of the Cambrian succession ofsouthwestern Sardinia and of r On the right: fossil content, sampling horizons and faunal ranges are shown for each level.

THE SEDIMENTARYSUCCESSION OF THE CAMPO The colour of the Campo Pisano limestones at PISANO FORI\4ATIONAT GUTTURU PAI-A, Gutturu Pala is mosdy red or light-red, only occa- sionally light-grey colours occur, while dark facies The Gutturu Pala localiry where the here report- occurs at the base of the lithostratigraphic unit. Near ed fauna comes from, is an abandoned sulphidi-ore the transition to the overlyirg Cabiva Formation, mine situated in the northern Iglesiente ereaabout 4 light-greenish/yellowish colours were observed. The km southeast of Fluminimaggiore (compare Text-fig. colours are bound to clay minerals, but show no dis- tinct distribudon paffern over the profile. After testi- I ). The late-diagenetic to epigenetic pre-Variscan mineralization (Boni 6( Balassone, 1994) has taken fyirg 13 samples (GP I to GP 14) for the claylquertz place within the massive carbonates of the San content, there is no very strong correlation benveen Giovanni Formation (Gonnesa Grorp), the overlyitg colour and clay amount or between occurrence of Campo Pisano Formation, interesting here, is effect- biota/biofacies and clay content, but it seemsto exist ed only locally at its base. The whdle sedimentary an indefinitely trend that the redder the rock colour successionat Guturru Pala (thickness is about 46 m) the higher the clay content. So, the colour may rep- resent.a of siliciclastic input, more is tectonically tilted, so that the stratification is more p.o*y. .the ,!r'r, or less vertical today. On the top^sediments of the Campo extensiveinvestigation is needed to verifr this indica- Pisano Formation, th. siliciclastic of the tion. Generally, the clay content of the samplesis less Cabiva Formation follow gradually. than l0o/o (averaged60/o). Much fewer is the quaru 386 o. ELICI(I, G,L. PILLOIA content: only in some horizons an amount of less PALAEONTOLOGY than lo/o was detected. X-ray investigations carried our in this section (Lecca et al., 198) revealed the To obtain pristine data, for fossil biodiversity esti- occurrence of quartz, chlorite, illite and Fe-oxides. mations solely thin-section investigations were used. Carbonate microfacies types of the Campo Pisano Chemical exiracted fossils were eiclusively used for Formation are generally very-rich in mud and include til(onomic work. mosdy wackeslones, iuboidinately mudstones and During our investigation the followitg groups of floatsiones. fu the siliciclasdc content, the carbonate organismJ were observed from the Gutturu Pala sec- facies rypes show no distinct vefticd Paffern of distri- tion: bution. many: trilobites,echinoderms, brachiopods, The most conspicuous feature visible in the field poriferids. is the nodular texture of large portions of the section. moderate: Lh"tt..lloriids, hyolithids. This diagenetic nodularity-is typical for many (not rare: hyolithelminthids,bradoriids, pelagi- for all) oTthe Campo Pisano seitions (Pl. 1, $g. t): ellids. of microfossils at The intensiry of the nodular texture seemsto {epend Generally, the preservation on a distinci minimum content of clay.So, a distinct Gutturu Pala is not uniform. Numerous specimens nodulariry was observed at Gutturu Pala ottl)' in lime- were silicified during diagenesis(.,g. the trilobites stones showin g a clay content more than about 60/o; and many of the brachiopods).Other ta>€ are Pre- in contrast, thEre are also (but rare) limestones with a served by original phosphate mineralory (...g. relatively high clay content (8.5o/o)which are hatdly hyolithelminthids, bridoriids, some brachiopods). nodulai (..E. at the transition to the overlyitg Most of the fossilsare disarticulated, but they do not Cabitza Foimation). Most of the limestones at showsigns of significantintensive or long-time ffans- Gutturu Pala are more or less wavy to bulbous by ^brokenport processes (tto abrasion,solution, Sorings).So, the end of origination of diagenetic nodules (by Pressure solu- individualsare not common until tion), which are separated from each other by accu- the Campo PisanoFormation. At the beginni$ of mulative seams of hon-soluble material. There is no the Campo Pisano Formation, the fauna is :l9Tly detectable correlation between the degree of nodular- dominatedby spongespicules (= fauna I , level GP I ; iry and the occurrence of carbonate microfacies tyPes Pl. l, fig. 2).-Ohly very seldom,some single echino- or the fossil content. derm phtes, trilobite iemainsor small brachiopods, Sedimentary features (e.g. erosive surfaces, cross- bradoriidsor hyolithelminthidsoccur. This basalpart bedding, grading, ripples, and bioturbation) were not of the Campo-Pisano Formation is overlain by an observ.?."V4t.tJat th; transition from the underlying assemblasewith a distinct hieher-(= amount on echino- massive limestones of the San Giovanni Formation derm "ttr trilobite remains fauna 2, level GP2- (Gonnesa Group) to the Campo Pisano Formation GP4A; Pl. 1, fig. 3).Alto brachiopods,ghan_celloriids (IglesiasGroup)-is rather distinct and sharp, the ffan- and hyolithids occur.Fau na 3 (levelGP4B-GP9) suc- siiion of the Campo Pisano Formation to the overly- ceeds,consisting qualitatively of the same content' ing Cabiva Formation is gradual over a Permanent but showing different quantitative ProPortions: inEreaseof the clay content-and a small alternation of echinodermJandtrilobites decrease significandy. The limestone and claystone within the last few decime- followi.g fauna 4 (level GPIO-GPI, A) is charac- ffes of the Campo Pisano Formation. terisediy ^ distinct increaseof spongeremains. So,

EXPIANATION OF PIATE I

Fig. I - TVoical nodular texture of the Campo Pisano Formation. Camera lid for scale. Fig.2 - Siiculite faciesin thin sectionat thi baseof the Campo PisanoFormation. Level GP I . S Fig.3 - gthinoderm faciesin thin section.Level GP3. Scalebar = 1 mm. Fis. 4 - Verv fossiliferoustrilobite faciesin thin section.Level GP14B. Scalebar = I mm. Fig.5 - foJt-r"y.d (a*0, only three preserved)chancelloriid scleritein thin section. Dark intern: vation of thet. remains.Leve-l GP3. Scalebar = I mm. Fig. 6 - Chancelloriidroserre in thin sectionshowing horizontal and verticd rays (8+1) and trilo =lmm. Fig.7 - Disarticulated echinoderm elementsin thin section. Middle of photograph: polygond th inlets (epispires).Level GP2. Scalebar = I mm. Fig. 8 - Oir"tri.ilr',.d echinoderm element. Samesample as for Pl. l, fig. 7. Note the preservatir GP2. Scalebar = I mm. O. ELICKI,G.L. HLLOLA, CAMBRIAN MICROFAUNAAND PALAEOECOLOGYC 388 O. ELICKI, G.L. PILLOIA

this fauna is quite similar to fauna 1 with the differ- ence that the trilobite content is higher in fauna 4. The overlyirg fauna 5 (level GP l4B)"shows a distinct decreaseof poriferids and an increaseof trilobites (Pl. l, fig. 4). Sb, the fauna 5 is similar to fauna 2, but shows fewer echinoderms and no chancelloriids and hyolithids. The youngest faunal association of the Campo Pisano Formation at Gutturu Pala (= fauna6, level GP l4C-GPl7) shows a distind decreaseof all faunal element and resemblesfau na 3 in its quantita- tive proportions. Summ arizing, the biofacies successionwithin the Campo Pisano Formation at Gutturu Pala can be Text-fig. 3 - Completed redraw of Eiffelia araniformis (Mis- interpreted as two similar pafterns (portions) from sarzhevski,1981), compare Pl. 2, fig. 14. Level fauni I via fau na 2 to faun i 3 ^nd, respectively,from GP2; max.diameter: 0.8 mm. fauna 4 via fauna 5 to fauna 6. If thise r,t.i.rsions indicate a trophic or a palaeoecolosicalchange or a changirg of liabitats ".i envirotrtti.nm wiil 6e dis- The remainsare badly preserved and mosdy represent cussed below. siliceoushexactinellids and secondaryphosphatised In many cases,systematic palaeontological investi- heteractinids.In somecases, it is not sureto d..ide if gation on ihe material attainid by chemical prepara- the remain representsa singleneedle or a broken part from a mor. tornplgx spicile..(Pl.2, fig.,,li).-This is tion or observed in thin sections is not possible. So, "monaxons" (Pl especiallythe casefor iome and cruci- the very rare hyolitfrids . 2){gr 3-4) a-re.only frag- "tetractins" mentary preserved becauseof the preparation proce- form (?stauractinsof lyssakidporifera), dures. The same is for the echinodeims: as riell in which occur upwardsin the section(level GP I 5). It thin sections (Pl. I , figr 5, 7 -8) as in result of the cannot be excludedthat someremains represent bro- chemical preparation (Pl. 2, Frg.11), the echinoderm ken partsof phobetractineelements as described from (eocrinoid) remains are represented by disarticulated the Campo PisanoFormation by Mostler (1985). (but not abraded) plates and so, uselessfor systemat- \Tithin the first 5 m of the profile (includirg the first ics. Also, the only r-emainof a stone-core of a pelagiel-"r..- poriferid maximum level),-only simple mbnar,ons, lid mollusc is relictic. Nevertheless, from oth.r cruciform tetractins and pentactins were found tions of the Campo Pisano Formation pelaeiellids (GPl-GP4). From the sameinterval the heteractinid have recendy been described (Elicki , Z00i- - all Eiffelia araniformis(Missarzhevski, 1981) has been observed specimens came from the same level near identified (Gxt-fig. 3; Pl. 2, figr 13-14). From the the base ofthe Campo Pisano Formation and belong secondporiferid maximum level, no specimenswere to only one speciei: Pekgielk subangulata (Thtel attained by chemicalpreparation. Aftir thin-section 1892). The problematic pelagiellid iemain from investigadon,the spiculeshere seemto be the same Gutturu Pda comes from a sffatigraphic comparable than those at the base of the formation. Further interval (GP2-level). chemicalprepared specimens come from the GP 15- A special point representsthe brachiopods. Many level (followirg the second poriferid marimum). specimens were attained by chemical preparation. Here, monil(ons and cruciform tetractins(= ?relicsof After first observation it seemsthat only n.ry few lin- phobetractins or ?stauractinsof lyssakid porifera) gulid and acrotretid (Pl. 2, figs 5-10) represent occur. ihe rather monotonous brachiopod fauna (friendly Unfortunately, the poriferid taxa from the personal communication by Michal Mergl, Cambrian of Saidiniapr.rbfirhed by Mostler (1985) PlzefilCzechi"). arenot indicatedto distinct localitiesor samplinglev- The main microfossil content and those which els. Only for his newly introduced' sp"ecies allowed a systematic approach are briefly Sardo sp o ngi a (= D ode caacti n eik) triradi ata, the locali- described "layer here. ry Gutturu Palaand a samplinghorizon named 150" is mendoned,but lacking any explanationor for. PoruruRrDS graphic ,hq position of that l"Vr!,gui11g th. pre- sent lnvestlgatlon,rro speclmensof thrs attrnrty were The occurrence of sponge spicules at Gutturu Pala observedfr6m this section. (ll. is verypatchy 1, fig. Zj Vi. ?, 4gr 13-22).Those remains predominate clearly at the lowermost part of CTnNCELLoRTTDS the Campo Pisano Formation (25o/o,levelGPI), but they decreasedramatically after some decimetres (to Chancelloriidsrepresent a problematicgroup of l-2o/o). A second marimum of the spicule content Cambrian spiny skelital elemerits.They werEregard- occurs at level GP lO/GP l4A interval- (about l5o/o). ed as potif.rids ('Walcott, 1920; Butterfielf, 6( CAMBRUN MICROFAUNAAND PAIAEOECOLOGYOF THE C, AIPO PISANO... 389

fragment (Text-fig. 5; Pl. 2, figt 24-25) t! nearly 3 -ri long and hasi circular crosi-section of about 0.3 mm. Th; outer surface is covered by more or lessreg- ular rine-like annulations in a distance of about 50- 90 U-."Th. relative tp(onomic uncertainty results from the bigger diameter compared to the holoryP.e (latter: 150 Ftl. All other charicteristics fit well with the original description.

Text-fig. 4 - Redraw of Chancelloria sardinica Mostler, 1985, comparePl. 2, fig. 23. Level GP5; mil(. diameter: rex'-ng5- t 1.3mm. f#; ?;,"ffi::{'*#afiTtftr',:aoi of tube:2.6 mm.

Nicholas, 1996) or together with halkieriids, wiwaxiids, sachitidsand siphonogonuchitids- as an PnoSLEMATICA own class or order named Coeloscleritophora more or (Bengtson6c Missarzhevski,1981). Followi.g A problematic remain, rePresentedby + Y.bl- to ?Aethol(opyltk j"tt.tC"t.tr( 1999), the latter is polyphylethic and the less ir^regular sphere is assigned Morris, 199"0 (P[. 2, fig. l2).- The chancelloriidsmay rePresenta non-PoriferangrouP adnata eot*"y a diameter of 0.5 lrllrl, numerous of hitherto uncertainaffinity. only specimen-has ronhd openings (about 30 and is slightly flat- At Gutturu Pala,chancelloriids were found only "b-ase". Fm) tened "t th. The sphereis hollow and siliceous in the lower third of the succession.The scleritesare in composition (secondiry diagenetic- gffect as in generallyvery rare ([essthan lo/o),orf in level GP2 manv associated brachiopods ind trilobites). The ih.y ^r, a titile enrich.d (gp to 5.o/o).The stone-core- shape-,size and di"-.ters of the openings preserved,phosphatised chancelloriid rays are often g.n.i"l to an "ffili"tion to the til(on mentioned above. by chemical procedures' 6oint disarticulated PreParation b..",rse of the bad original mineral- chancelloriidrosittes can be observed Preservation,'the but sometimes opy and the internal structures are not preseryed. So, (Pl. Pl. 2, fig. 23). "envelope" 1, figs 5-6; tliir siliceous may represent only the single rays' fw9 Besidis the usuallydisarticulated replaced outer wall of Aetholicopalla adnntA. The fact preserved taxa were observed: more complex that (in contrast to the specimens described by Mo.stler,1985 (Text-fig.4; Pl. Chanceltoriisardiiica Conway Morris, 1990 and by.Elicki, 1993),rlt. P,re- 1965 (Allonig 2, fig. 23) and Allonia Dord 6( Reid, sent speclmen is hollow can be interpreted in that chancel- sp.)."'Amore detaileddetermination of the wav th", the primary mineralogy was rather calcare- totiia remains from Gutturu Pala is not reasonable o,ri and diaeeneticaliy dissolveJ'than phosphatic - a becauseof the morphological variabiliy of those question *Ki.h, hitherto, could nol been surely rarenessof the findings. remainsassociated wilh the ihferred. A decision of the Present specimen to the Mostler ( I 985) describedhis new erectedspecies problematic structu re Archaeobidesis unlikely "layer similar "base" Chancelloriasardinica from 146" (but, again, because^ofthe lack of the flattened and of dif- givesno explanationfor the position of that laygr). fering surface sffuctures in the latter. The biological Fot the fuither chancelloriidsmentioned by that affini-'ry of Aetholicolalk is sdll unclear. Surely, they author, no localiry is indicated. do not represent f'ree-livitg metazoans' b,tl-aybe resting bodies for storage of ProPagules (Conway HvoIITHELMINTHIDS Morris, 1990).

Hyolithelminthids are widely accePted.as Pgly: AnrHnoPoDS cha.tl-worm tubes.Typically, they are small conical phosphatictubes with a more or lessirregular curved Ostracods from the Sardinian Cambrian were apicai area.Toward th9 aPerture' .tt.y are elongated reported up to now only by Mostler (1985) dg. to Cambrian "ird more straighten.The crosssec{ql Tay be circu- obr.*ations in thin sections of Early (hyolithellids) (torellellids). archaeocyathidlimestones. He described ostracodsas lar or elliptical "occasional At Gutturu Pala, rare and bad preservedspeci- occurri.g, strongly recrystallized' not mensof this fossilgrouP were found exclusivelyat.the nearer definable, r-ill shellst.'Also itt the Campo base(GPl-level). The-specimens were identified as Pisano Formation ostracodsseem to be rather rare: in the lowest level (GP t only few specimensof Hyotithelluscf . filiformis Bengtsotr,1990. The longest only I 390 O. ELICIQ, G,L. HLLOIA

localities of S\f Sardinia, include Protolenus (Protolenus)pisidianus Dean (in Dean 6c Ozgtil, 1994).The vertical rangeof this latter Partiallyover- laps the succeeding Acadoparad.oxidesmureroensis Zoneof the basalCFZ trilobite assemblage.The sPec- imens tentatively assignedin this p"p.t t"oCkuigellus belong to the CP I fairna (seediscusiion below). The CPz fauna clearlyshows a more diversifiedtrilobite assemblage, includitg the above mentioned AcadoparZdoxides mireroensis and Protolenus rext-ng6- (Protilenus) pisidianus at the base, while orycto- !{,(,"';!ff#,:!:::;#,t'lo;,;?T!"ff', :'f:lx cephalidshave been found exclusivelyin the middle GPI; mil(. diameter:2.9mm. p"^tt of this formation, associated with Acadoparadoxidesmureroensis, Calodiscus foueolatus Howell, 1935, and dorypygiids. In the uPPer Part asnostiids(Peronopsis, Prychignostt ts, Dip kXnostus)are this fossil group'-B..iuse could be proved (Text-fiB. 6; Pl. 2, figr l-2). of th6 very charact6ristic and alsociatedwith ParadoxidesIp., Pardailhinia hispida (Thoral 1935), Corynexochusdelagei (Miquel 1202), s_tionglysignificant -needed morphological features, it is justi- fiable"and to estiblishl new til(on on species sp., Dorypyge sp:, Ctenocephalus level(see below). (Ctenocephalus)sp., and Dawsonit. Thilobites from the Campo Pisano Formation The specimensobtained by chemicalpreParadon_ havereceived little affention uhtil now. Despitetheir are usuafly tiny and more adapted to t6e itudy of abundanceas tests detectable both in the field and in small trilobit.t "t agnosdds"td eodiscidsor imma- thin sections(since Bornemann, 1886), systematical- ture develop-e.tt"I stages of larger individuals. aboit largertrilobite ly useful remains are difficult to collect an4 to P{e--the Conseq.r.ttfy, the inform"ations p"r.. Rasetti (1972) describedfew til€ from remains from the Campo Pisano Formation at locdities l2r and l3r at Corovau (northern slopeof Gutturu Pala are missing. In this paper the occur- According rence of Condylopy7tantiquA n. tp.: Sf e$. Marganaimountain range,?o.-"snovas). UiaEoltus to that author, the faunl originates from the uPPer pompechiiand representativesof Aconthetnae ls Campo PisanoFormation (cohsideredat that time as repottedfor the first time from the SardinianMiddle uppermost Gonnesa Formation) and includes Cimbrian; in addition, the Gutturu P{a faunaallows Agrarlos sp.,Corynexocltus ktus Rasetti, 1972, C. sAr- . ia."itfy Dawsoniaes D. bohemica(3n^j4r, 1950). rtut Rasetti, 1972 (both junior synonyms of C. Due to the pauciryand preservationof the arthro- delagei), Pard.ailhania_hispida (Thoral, 1935), pod taxa, desciiption is reitricted to the best repre- Pero-nopsissp. and Ptychagn6stussp. Later orl, sParse sentedand preservedspecies. inforniation was giv-enby Pillola, accompaniedby illustrationsof the"'b.tt pt'.servedand biosiratigraph- ClassOsrnncoDA Latreille, 1806 icallv sienificanttil€ from the so cdled CPI and CPz Order BnnooRIIDARaymond, 1935 faunas,"without systematicdescriptions (Pillola, Family-Genus HIppoNICHARIoNIDAE Sylvester-Bradley, 196l l99l; 1994;Loi et'a|., 1995;Pillol a-et al., 2002 and HIppoNICHARIoNMaffhew, 1886 cited references). - Thilobitesof the CPI fauna, recognizedin few Ttpt species HiPPonicharioneos Matthew, 1886.

EXPIANATION OF PIATE 2

FigsI,2 Hipponicharion ichnusurnn. sp.; samespecimen in both pictures, stereo-photograPhwith u paif of marsin, holorype.Level GPl. Scde bars= 0.5 mm. Figs3,4 Unassiened"hyolithrerinains. Levels GP8 (fie. 3) and GP2 (fig. 4). Scalebars = 0.5 mm. Figs5-8 Unassilned"itott.tid brachiopods.Fig.6 is-detailof (5). LevelGP15 (figs 5-6,8), GP9 (fi Fig.9, l0 Unassilnedlineulellid brachiopodsin phosphadcpreservadon. Level GPl. Scde bars = 0.2 Fig.11 - DisartiZulatedEchinoderm remain. Level GP2. Scile bar = 0.2 mm. Fig. 12 ?Aetholicopalkadnata Conway Morris, 1990. Level GP2. Scalebar = 0.2 mm. Fig.13 Smdl specimenof Eiffelia ariniformzs(Missarzhevski, 1981). Level GP2. Scalebar = 0.2 m Fig. 14 Bad pt.r.*ed?Eiftfi7 araniforinzs(Missarzhevski, l98l). Level GP4. Scalebar = 0.5 mm. Figs15-22 Unasilgnedsponft spicules.Level GP15 (figs 15-19),level GPI (figs 20-22). Scalebars = ( mm (fiet 15, 17,18). Fig.23 Chancelloriasardinica Mostler, 1985. Level GP5. Scalebar = 0.5 mm. Figs24-25 Hyotithellus cf.fiWrrnis Bengtsotr,1990. (25) is detail of (24). Note the ring-like ornamen Scalebars - 0.5 mm (fig. 24\,0.2 mm (fig. 25). O. ELICKI, G.L. HLLOLA, CAMBRI.ANMICROFAUNA AND PALAEOECOLOGYC 392 O,ELICKI, G.L,PILLOIA

HlppoNIcHAzuoNICHNUSUM n. sp. the Early-Mid Cambrian boundary in S_ardiniais sit- Pl. 2, figt l-2; Text-fig. 6 uated ,err.tal metres above the base of the Campo Pisano Formation, that means that H. ichnusumis lateToyonian (= late Bilbilian) in age. 2003Hipponicharion sp.- Eucruet al., Pl.5, ft1.2- P.33, Th; palaeobioseosraphicdisiribution of the genusHipponichaion TvIinhew, 1886 was discussed Holotype- Right valve;FG 53611(Pl. 2, frg- 1). 5y Gozafo 6( Hinz-schallreuter (2002)' comitg to the result that the genus is restricted to western Repositor! Geologi."l _ Institute, Freiberg Gondwana,Avalonia,.'and Baltica and so, rypical for Universiry archivenumber 536. the AcadobalticProvince sensu Sdzuy (1972).

Deriuation of name After its occurrence in ClassTTIoBITA \falch, l77I Sardinia(historital name: Ichnusa). Order AcNosrIDA Kobayashi,1935 SuborderAcNosrINA Salter,1864 and locality - B{"1 Campo Pisano Typt horizon Superfamily^ CoNoYLoPYGoID_EA Raymond, 19| 3 Formation (first 0.5 m; l6vel GPI), Guituru Pala Family CoNDYLoPYGIDAERaymgnd, 1913 mine, about 4 km SE of Fluminimaggiore (S\f GenusCoNo)aLoPYGg, Hawle & Corda, 1847 Sardinia). - Tvpe'L species' Battus rex Barrande, 1846 Material - Three specimens. (by monotypy),Bohemia.

Occurrence Late Early Cambrian (latest CoNoYLoPYGEANTIQUA n. sp. Toyonian = latest Bilbilian), Campo Pisano Pl. 3, figt l-12 Formation. 2003Condylopyge sP.- Ellcruet al., P. 29,PI. 3, figs3-7. - large (maximum lengtht Diagnosis C,"t?p,ace l.S mm; mililmum hei[ht: 2.1-mm) and sub-triangular Holotype- Cephalon,DSTCP 23155,Pl. 3, frg.5. in lateralview. Dorsal margin straight.Ventral, ante- - Terra, rior qnd posterior margins gTly fragmentary Pt.- Repository Dipartimento di Scienzedella served,but if visible without ornamentation. Caglilri University,acronym DSTCP 23. Lateralsurface smooth. V.ry long anteriorand Poste- "ancieflt". rior crest-likelobes parallel to sub-parallelto_ the mar- Deriuationof name-From the Latin gin and reachingtteir to the dorsalmargin. Ventr"lly' ih. lobesapproich eachother very, but they do not Typt horizon and locality pampo. Pisano comein touih. Anterior lobe nearlystraight' posteri- Formition, level GP9, 24.7 m above the base; or lobeslightly curved.Mid-dorsally a circularnode Gutturu Pala section, Fluminimaggiore, S\f is prominentlydeveloped. Sardinia.

Discussion The new speciesdiffers from the Material Level GP9: one poorly preserved other known hipponicharioniidsas follows: from H. enrolledspecimen, I I cephalaand 4 pygidia,DSTCP Hinz- eosMatthew 1886, H. midahensisGozalo 6( 23001-0i, 23016-17, 2lozg-31 and- 23155; level geyeri 1993 Schallreuter,2002, H. Hinz-schallreuter, GP14: 4 cephalaand 2 pygidia, DSTCP 23065-69 2002 it and H. elickii Gozalo& Hinz-Schallreuter' and z3l54.'Additional qpe-imensfrom loose block more sub-triangula-r o{ rlt. differsclearly by the qhap. GPLB: 19 cephalaand 11 pygidia, DSTCP 23082- the sjzeand shapeof ihe lobes,"td b), carapace,by 100, 23ll 8-[zt, 23135-137, 23141-144. the b...tttetc. of a prominent mid-dorsalnode. A further difference to H. geyeri Hinz-Schallreuter, occurrence Early Middle cambrian, campo 1993 is the lack of pits. Fro-rnl/. hispanicumGozalo PisanoFormation. & Hinz-Schdlreuter, 2002 the new speciesmainly differs in the shapeof the caraPaceand in the Pres- Diagnosis A CondyloPygtspecies crescentiform enceof a prominint mid-dorsalnode. relativelysmall anterior lobe, strong.sp.ines The biostratigraphiclevel of the finding horizon shaped, and at the_posterolateral.edges, of Hipponicharionilhnusum results from the position otr ?h. occipital-ring -m field and well-develgpgdborders, in 0-0.i abovethe San Giovanni Formation (Gon- long ^..ih"lonpreglaEellar Pygidial oil slightly nesa Group). The top of the latter was biostrati- botfi and pygidium. faint fur- graphically^defined bi Perej6net al. (2000) on ,l]. taperingbachvards, biaring the trale of tree risen E"rir of archaeocyathias Toyonian 2-3 (re9pe_ctively rows and a poorly marked keel, occasio"4ly mid to late Bilbilian, latestEarly Cambrian).On the into a faint node; obsoleteor faint impressedmedial other hand,after Loi etAl. (1995),based on trilobites, furrow posteriorto the pygidial axis. CAMBRUN MICROFAUNAAND PALAEOECOLOGYOF THE C/ MPO PISANO,,, 393

Description Cephalon with parallel to gently CondylryStrp. Alrraro&.Yizcalno, 2000, from south- roundedsides, anterior margin -both well rounded.Narrow ern France. border and border furrow, broading and shal- The Sardinianspecies is very similar to C. globosa lowing in correspondenceof anterolateralregions (Pl. (Illine, l9l6), which differsfrom C. antiqua in hav-' 3, fig. 6); short and strong paired lateral spines, ine ishorter preglabellarfield and narrofrer border, extendirg baclnvardsfrom genal region. Preglabellar thE posteriorfobJ ir slightly wider than long; C. gh- field long (sag.),about ll2 of the anteriorlobe, sepa- bosaalso differs in having weakeroccipitd and pos- rated from a raised anterior border by a well- terolateralspines.- impressed border furrow. Axial furrows deep and A close speciesis representedbv Condylopyge sharp. Glabellaconvex, futgiform in shape,as long_ 'Warwickshire,amitina Rushton, 1966,^from the Purley Sh-alJs, (sag.)as 70-8 0o/o of the cranidiallength, consisting of which mainly differs from C. antiqua two lobesseparated by deep,gently anteriorlyarcuate by the absenceof the medial furrow, by the unfur- ffansversefurrow. Anterior lobe crescentiform,sagit- rowed and more rounded pygidial a)ris,'andslightly by the proportionsand shapeof the remaini.g parts tal length about half the width, extendedposterolat- 'btaeru.r" erally,occupyi.g 28-30o/oof the cranidiallength (lit- of the carapace.Condylo?y7t Fletcher,1972, tle less in immature stages).Prominent posterior from S\f Avalon/Newfoiridland, shows closesimi- glabellarlobe, sub-ellipticalto rounded sub-quadrate larities in the ^rh"'ort.rpveidial features but the cephalondif- i'n outline, slightly longer (sag.)than wide (tr.) with fers in having preglabellarfield "ttd cephalic genal obsolete median ridge toward posterior, and very borderand loneerspines at the cornersand on possess faint median tubercle at posterior end; narrow and the occipital ririg. Cond.ylopyget/ortnszi short- pysidal pair of low occipital lobe of about equal width (tr,) as the er oris ind a mot6-riarkedkeel, and a ploiril pvsidium; in addition, tlie bor- anteriorglabellar lobe, composed of a pair of swollen furrowsin the marsin are relativelynar- basallobEs and a middle portion mostly represented i.r furrow and the bdid"er row and showsrather uniform"'width, both itt the by the baseof a well developedposterior occipital cephalonand in the pveidium (Lifidn & Gozalo, spine.Genal region narrow around the anteriorlobe, 1986:pl. 1, figs 1-8).The-occurrence of a pairof lat- broadensadjacent to posteriorlobe. eral fuirowr "itoss the pygidial pleural field allowsto Unreleasedthoracic segment (Pl. 3, fig.7) showsa distinguishC. regiafrom the Sardiniantaxa. well-developedmedian node on the axial ring. CJmparisons-of C. antiqua with the only aqd Pygidium sub-ovatein outline, with well-developed incompli,tecranidium of Coidyhrygt sp. from iouth- border. Axis convex, gently tapering poiterolateral ern Montagne Noire, France (Alvaro 6( Yizcaioo, (sag.) of whole bachvards, occupyitg 55 to 75o/o 2000, pl. ll fig. I is delicate;however, the French ll2 ) pygidial sagittallength and transversally,about of species^shows Zlearly narrower cheeksclose to the ih." marim"umwidih of the pygidial shield, showing ahterior lobe and rhott.t (sag.) preglabellarfield, more or lessmarked trace of tree pairs, shallowitg which sussestmore affinitieswi-th C. rZx.Similar dis- bachvards furrows, impressedonly at adaxial ends tinctiv. fEi't,rresare present in the CondytoPygtspecies (ll. 3, fig. 8) and a poorly.-"Tkgd keel,occasionally figured by Geyer ( 1988) from Morocco. CondyloPtgt nsen lnto a faint node. Faint shallowmedian furrow eli from the Moroccan Anti-Atlas shows a relatively on posteriormargin. Borderwide, particularlydevel- shorterpreglabellar region and faint cephalicspines, oped and shallowerin correspondenceof the postero- lonser pveiaid axis aid lessrounded outline of the lateral regions;gutter-like border furrow, shallower cepf,ali;;d pygidial shields. ancand wlcerwider at posterolaterdposterolateral Poruons.portions. lvreasuremeMeasurements of holorype: sagittal cranidial length about 2 mm. Family uncertgin GenusSrcngncNosrus Snajdr, 1957 Discussion- Condylopygeantiqul n. sp. differs sig- nificantly from most describedspecies of Condylopyge Typtspecies - SkryjagnostuspomPechji Snajdr, 1957; in havirg a medialfurrow posteriorto the pygidiala:ris Bohemia. and a posteriorlytapering rachis, which allow a clear distinction againstCondylopyge rexBanande, 1846, C. SrcnqecNosrUSaff. poMpECKJISnajdr, 1957 carinataVestergird 1936,and similar species. Pl. 3, figr 13-20 Closely related Condyhpygtspecies, showing a faintly furrowedand not posteriorlyexpanded pygidi- Material- GP9,4 cephalaand 2 pygidia,DSTCP urn, includeC. cruzensisLifuin& Gozalo,1986 (name 23010,23018,23035-39; GPI l, I cephalonand I modified in cucensisby Geye\ 1998),from Spain,C. pygidiurrr,DSTCP 23051-52;GPLB, 3 cephalaand eli Geyer, 1998, from Morocco, C. gtobosi (llling, 3 pygidia,DSTCP 23131-34,23138-40; totallv 8 1916)and C. amitina Rushton,1966, from England, cephalaand 6 pygidia. C. regia (Sjcigren, 1872), from the Eccaparadoxid.es "etaerus" insularis Zone of Sweden, Condylopyge Description Semi-elliptical,stronglv voulted Fletcher,1972, from SVt Avalon/Newfoundland,and cranidium,slightly wider (tr.) than long (sag.),very 394 O, ELICIQ, G.L. PILLOIA frintlv defined or completely effaced tapering for- ellipticd than sub-circularin shape. *ards glabella.Posterioi end of the glabeflaincfined, th. genus Skryjagnostus6 known from the with o"ccasionaloccurrence of e eitremely narrow Middl. i"-brian Firidoxides -orpusillus Zone of Czech (tr.) (Pl. fig. 16). Republic and Pedinocephalus TbxotisZone from (sag.)and Jotg occipitd ring 3, Presenceof a narrow doublure. noithwesternSiberia. Pygidid shape nearly'all idgntical to the cephalotr: litde'irot. voulted in directions, with defined SuborderEoolscINA Kobayashi,1935 rachis, especiallyclose to the tiny articulated half Family EoolscIDAERaymond, l9I3 ring. Rachis,o.i,rpying about 85-iOo/oof the pygidi- GenusDn''urrsoNlA Hartt in Dewsoo, 1868 al length (sag.),usually not clearlydefined at the Pos- teriorlnd and expandedand slightly more elevatedat Ttpt species- Microdiscusda.wsoni Hartt in Dawson, g6g. middle lenesh.t' ".., of a fain"t titto* border and I doublure J"t be observed (Pl. 3, figt 13, I7). Cephalonlarger: about I mm long (t"g). De''urrsoNlABoHEMICA (Snajdr, I 950) Pl. 3, figt 2l-26 Discussion- For many of the remains, the assign- nov. gen. nov. subsp. ment is particularlydifficilt due to the absenceof dit- 1950 /cuhodiscus bohemicusbohernicus SNeJon,p.201,pl.1,$g. 1;pl. 2,figsl-6, Pl.3,figsl-2. s and to the q?e of preseruation- not - tinctive h"t.tt 1953 Aculeodisiusbohernicus $najdr $ncr-uR, p. 490-491. only to the to(on but also for e certain distinction 1957 Acuhodiscusbohemicus $najdr -$Ne;oR, P. 240. between cephala and pygidia! Due to the peculiar 1958 Acuhodiscusbohemicus Snaidr - SNnlon, p. 46, Pl. 1, figs l- morpholog;r of this t*ioi, only few details can be 22. - & Bnsn, p. 81, to"ih. orieinal diasnosis and description 1970 Acuhodiscusbohemicus Snaidr Honxf addid pl. 1, fig. 10. (Snajdr, 223, in Eriglish) addidonal gl7. 1957,p. -and.on 1990 bo*rofio bohernica($najdr) - SNnlpR,P. 86, text-fig. illusirationsgiven by the samt author (Snajdr,1958: 1992 Dawsonia bohemica(Snqdr) - Fnrre & KonpULE, P. 49, pl. 6, figt 4-t, 10, 12-20).The SardinianS[ryjagnostus fig.2. shows i clearly smaller articulatedpygidial half-ring v1995 {f . Dawsonia- Ini egal. 68, Pl. 4, fig. 9. (Sgejdr) - 2. (transversdlyless than one third of the pygidium at 1996 Dawsonia bohemica KgnouLE, P. 43, fig. 2000 Dawsonia bohemica (Snajdr) - Arvano & VtzcelNo, P. the samel*.1) and poorly defined anterior border. In 279, pl. l, figt 2-6. conffast, the pygidial rachisis usually better defined. 2003 Dawionia bo''hemica(Snajdr) - EucKr et Al., P. 29, pl. 3, In addition, 5;;h, cephalon and pygidium are more figs l-2.

E)(PIAI{ATION OF PIATE 3

All illustratedspecimens (excepred fig. 38) are housedin the Dipartimento di Scienzedella T Pillola Collectiin (DSTCP), t'oflo*etr by ihe specimennumber ind the acronym GP (for Gu The abbrevationtb is assignedto the materiaforiginated from a looseblock. Magnification : Figs'-'2 i;ilil.;;ftJHifrillBsT3fltrt',:#fni?rTf,i;"??,',i 23083',GPLB;'5) Incomplete iephalon, holorype., DSTCP. 23155, GP9; !)^Lqa. il:qryp zlogo',GPLB; Z) Imm"ttrt pygidi,tmwith uirieleasedthoracic segment DSTCP. 23154: DSTCP.23069, GPl4;9) Intbirpletepygidium DSTCP. 23094, GPLB;.lq) Pygidium D Incompletepygidium DSTCP. 23097,GFLB; L2) Lerge fragmentary cephalon, interolate -2023001, GP9. '.*Uil'.+Llil"ii{'ES8 Figs'3 "glw'#i# oit{,'{yi+fiil*,lflii ;: lL If : g'i6:t'd'#3dtr;iil'ffi'#s€?'bilil'trlil:" -26Bfl r;i',!il;^;'I3J?.ff Figs2' ?#;#:l!:K:#,1siH,1t;33:)63?Jfi?lt';:'"*iHfi r?lli?'??J3; view,bSfCp. 23f06, GPLB; 24) Pygidium,DSTCP. 23108, GPLB;-25) Dfitoned pygid Damaeedpveidium, DSTCP. 23023, GP9. Figs27-34i;y;f'&r{yrf'r*:xf .?l,Lill"lrJfei')fr ,{r:'a'r'r?ifr !1;?"?; Damasedimmature cranidium, DSTCP. 23027,GP9; 32) Crerridium,DSTCP. 23060, ( cranidlum,DSTCP.23059, GPll;34) Thansitorypygidium, DSTCP. 23114, GPLB. Fig;r35-37 Chutgeltus?l. sp.35) Cranidium,larvd I!3&e:?!1.9? 23152,GP2; 36) Cr 3D licomplete-immaturecranidium, DSTCP. 23045, GP2. Fig.38 - Unassignedcorynexochiid, DSTCP- 23063, GPI I (not describedin the text Fi;.39 - Dorypieid, DSTCP.23042, GP9 (not describedin the text). Fi;. 40 - ?Adauliid,DSTCP. 231 16, GPLB (not describedin the rext). Fi;. 4l - Ellipsocephalidsen. er sp.indet., DSTCP. 23122, GPLB (not describedin t FiE 42 - Elliirsoc.ihdid [en. et sir.inder., DSTCP. 23050, GP4 (not describedin tht O. ELICIQ, G.L. HLLOIA, CAIVIBRTANMICROFAUNA AND PATAEOECOLOGYC 396 O. ELICKI, G.L.PILLOIA

Material - LevelGP9: 5 cephala,DSTCP 23009, Order ConvNEXocHIDAKobavashi, 1935 23019-21,23040 and 5 pygidia,DSTCP 23011-12, Family ConntEXocHIDAEAngelin, 1854 23022-23, 23041; levef-Gpt t' 3 cephala and 2 SubfamilyConvNEXocHINAE Ahgeli.t -1854 pygidia, 23053-57; level GP14: one cephalgl GenusConYNEXocHUS Angelln, 1854 - ijS:fCP 23070;loose block GPLB: 11 cephalaand 3 U?t species Corynexocltusspinuhlus Angelin, 1854. pygidia, 23101-23109,23110, 23145, 23147-51; iot-"lly 20 cephalaand l0 pygidia. connrooc;i5,Tf#jf is".t,re 05

Description Coarsely granulose semicircular 1905 Corynexochusfukgei n. sP.Mtqunl , P. 481,Pl. 15,figs 4- cranidium, slenderelevated conical glabella,occupy- 4a,b. ine 650/oof the cranidiallength (ZOo/oandless in early l9L6 Corynexochusfukgei Miquel - \(nrcorT, p. 317,pl. 55, stages)and about 30=o/oof maximum width of figs3-3a. er6*th - ih. shieldit the base.Glabellar segmentation effact- 1934 Corynt*ochusdekgei Miquef LnxE,p. l81t - Mi{uel - THoner-,p. 46,pl. 3, figs traceof the occipital furrow, rePresent- t935 Coiynexochusfuligei ed, very faint 6-7. paft of narrow incisions,placed low on sides - ed bv a 1936 Corynexochusdekgei Miquel l.ttltR, P. 27.. of the eiabella,angling back beneathmedian srrong t96t Coiynexochuscf.trekgei Miquel - Sozw,334 (616),pl. *.1 developeacranidial spine arising from Ll 22, fig.6. and - (Pl. figr 2l-22). Narrow to absent depre^st:d 1963 Coryiexochusdekgei Miquel \7nrrEn, p.-369. 3, - Sozuv,104, pL.5, figs the anterior tips of the ? 1968Corynexochus aff.trekgei Miquel preglabella"rfield, flanked by lr-12. genae.Border furrow well impressed - itro'-telyconvex 1972 Corynexochussardous RnsErrl,P. 63, pl. 18,figs I -17: thoro"ugh,post.tlot borderfurrow shallower.Anterior 1973 Corynexochusdekgeilvliquel - CoURTESSoLE,P. 136' pl. border".r.3..ntiform, wide (sag.)as 20-25o/oof the 3, figs8- 1 8; pl. 25,fig. 6; Pl.27 , Fte.7 . - SHERGoLD& Spzw, cranidialsagittal length, narrow-ingtowards the g."4 ? 1984 Corlnexochuicf. dek[ei Miquel g or lessdeveloPed . P:74, Pl'l' fig29' ansles,rypiaally bearin 16-20more ? 1984 Corynixochusif. sardoersRasetti - SHencolo 6c Sozuv, t"Jid iniisioni defining sub-redangular-,,scrobicu- p.i4, pl.2, figslo. lae". Posteriorborder *id.tts v 1995 Corynuochusirdo^ Rasetti- Lot etql., P.-6^8, Posterolat.r"lly. - Pl.+, 19.7. Pygidiumsemicircular in test clearlyshow- v 1995Cofi,nexochus htus Rasetti Lol e,tal., p.-68: Pl. 4:.fig. 2. . *"P., - VtzcelNo, granulations (Pl. 3, figt 24-26); convex 2000 Corynexochusdekgei Miquel Arvnno 6c P. rns coarse 280,pl. l, figt 7 -ll. .oii."l rachfsdelimited by deep a:iial furrows, reach- about ing the posterior border furrbw, occuPyitg Marcial - Level GP9, 6 cranidia and 2 pygidia, rings and 3do/o(tt i of the pygidial width. Five orial DSTCP 23013-14,23024-27, 23043-44; level GPI l, sharp furrows. terminal piece, *e-ll seParatedby 5 cranidiaand 1 pygidium,DSTCP 23058-62,23064; Pleural t.giott convex, with 4-5 broad pleural ribs level GP14, 3'i{anidia, DSTCP 23072-74; level and ,-6 J..p interpleural furrows, shallow border GPLB, 2 cranidiaand 5 pygidia DSTCP 23Ill-L4, (Pl. furrow and fl"t to gently convex border 3, figt 2380;, nl59-60; totally i6-.t"tidia and 8 pygidia. 24-26). Description Exhaustivedescription of this til€ Discussion- The concePtand the historical con- has been made by severalauthors; more recendy, troversy about the genus Dawsonia (Hartt in Al.'ato &. Yizcaino (2000) had the possibiliryto re- Dawson, 1868) hasbe."" recentlydiscussed by Alrr-aro examine the Miquel and Courtessolematerial and 6{ Vizcaino (2000); contemporaneously,the first add the first desiription of the hypostoqe and ros- occurrenceof D. bohemicaoutside Bohemia and trum. The new materialfrom Guftirru Palais mostly comparisonswith congenericspecies have been made representedby immature cranidia and tiqlr P)'gi4?: by' the sameauthors. the smallestimmature cranidium (Pl. 3, figt 27-28) In S\f Sardinia,in addition to the Gutturu Pala showsa narrow (tr.) parallel sided posterior Part of section levels GP9, GPl l and GPl4, Dawsonia the glabella,which stiongly e*pandsforwards, with bohemicaoccurs in the CP2 assemblageof the deeD"eninedorsal furrows-into I pair of pits, placed Villaggio Norma section at Monte San Giovanni iust'belo# the largest(tt ) portion-of the glabell"(fl. (Goririesaeasternmost end of the ,,Cabitzasyncline") ?, fis, 2n S[ehily largef cranidia clearly show the and in rare pebbles originariqg ft9- the Campo palpEbraliobe i'nd'auefi faint ocularridgl (Pl. 3, fig. PisanoFormaiion, within the Caradocianconglom- lZ, risht side);at this stagethe palpebrallobe is about eratesof the Monte Argentu Formation, north of one tKird of the slabellar-length(exsag.), the anterior . D. bohemiiaappears in Bohemiain the tip is closeto the"dorsalfurrJw and tlie Posteriorend Paradoxidespusillus Zone and in southern Franceat fir ^way from the glabella and from the posterior the lower occurrence of Badulesia tenerA in the margin of ,h. craniJium. Unfortunately, the largest Coulouma Formation, suggesting an early cr"trldium is poorly preserved(Pl. 3,.fig.??) -an.d,as Caesaraugustianage. for smaller specimens,nothing can bJadded about segmentationor peculiaritiesoF the surfaceof the test CAMBRAN MICROFAUNAAND PAIAEOECOLOGYOF THE CA]IPO PISANO.,. 397

(minute quartz crystalssubstituted the carapace).The Occurrence Corynexochusdelagei occurs in availabl. Fygidia also suffer of the above-ci^tedefface- Montagne Noire (France) since the base of the ment and display more or less segmented rachis, Coulouma Formation up to the Al-A2 levels of pleural field s-ooth or faintly furro#ed and margin-"(Pl. Courtessole,in Spain (Asturias and Cantabrian "l futtow usu"lly obsolete. The figured specimen Mountain Range) and Sultan D"g (Taurus 3, fig. 34) is about I m1n large (ir.) grd refresents a Mountains, Tirrkey); in southwestern Sardinia, transitory pygidium with incilient hints of releaseof Corynexochusdelagei ranges from the middle Campo lts antenor segments. PisanoFormation and is also known from the basal levelsof the CabitzaFormation, a long rangedistrib- Discussion The high morphological variabiliry ution which coversthe upper Leonian and most of during morphogenetic development 6f Corynexochus the Caesaraugustan(includirg the Pardailhaniahisp- dekgii, the ii"giostic characters consid.t.d"by sever- ida beds of the CPz- ,,assemblage", th. al authors and the pt:servation, represent the rype .of Solenopleuropsis(Manubtesia) ribeiroi beJs of the main source of uncertainty for a safe assignment of CAB I and the Eccaparadoxides pusillus specimens- to this taxa. Eccaparadoxidesmediteuanius of the CngZ levels. The shape of the glabella changes considerably from sub-parallel sided up to the middle length to SubfamilyAcoNTHEINAE'Westergird, 1950 more or lessclavate both during growth but also with GenusCt wIcELLUSGeyer, 1994 distortion/compression during diagenesis (see the imprint of defoimation in terrifenoulmaterial figured Typtspecies - Clauigellusannulus Geyer, 1994. by Courtessol. ( 1972, pl. 3). In tiny specimens,such as the cranidium figured by Rasetti (1972, pl. 18, fig. CmvIcELLUS?n. sp. 3) the glabella is irore expanded anteriorly than i"n Pl. 3, figs35-37 larger specimens and resemblesclosely an incomplete cranidium assignedto C. ktus (Rasetti, 1972, pl. 18, 2003Ckuigellus sp. - Ellcru etal., p.29, pl. 3, fig. 8. fig. 2l). In otlier specimens, such as the cranidia fig- - uled by I oi et Al. (tgg5, pl. 4, figs 2,7) the glabelllr Material 3 immature incomplete cranidia from shapeof the specjgen,on h,isfi,g. Z sugges^ts.thEassign- level GPz. ment to latus while the obsolescence of the furrows suggestsimilarities with those assisnedto sardous.The Description Club-shaped unfurrowed glabella r"ii5 between the width (tr.) of Li and the maximum gendy expandirg forward from SO and constricted at width (tr.) of the glabella has received an elevated tax- the level of Ll, without preglabellarfield and anteri- onomic value in discriminating til€ of dekgei affini- or border, elevatedabove fixigenae; large sub-triangu- ties; however, as pointed out 6y Akaro U"Yizcaino lar to well rounded occipital ring, slightly larger (tr.) (2000, p. 280), this ratio ranges from 33o/oto 50o/o. than Ll, bearing a low backward and upward direct- The ociurrence of the occipital spine and the shape of ed robust occipital spine. Sub-triangular, well-round- the occipital ring represetttid furiher maffer of debate, ed fixigenae, gently convex in all directions. Palpebral but again, the variabiliry,observed on figured lobes crescentiform, aborially inclined, occupyi.g lpeci- mens seemsmostly related to different preservation. (exsag.)33o/o rc 42o/oof the glabellar length (excluding About pygidial'features, we can noticf that there are the occipital ring), placed about at mid-len$h of the sffong diffelinces in segmentation comparing internal glabella; faint and narrow ocular ridges, slightly moulds and externd surfaces(see Rasetti, 1972, pl. 18, obliquely directed to the largest portion of the glabella. figr 6-9, l?-!4, 16) and, substandally,there a5eno rig- Anterior branches of facial suture short (exsag.) ni-fic"ttt differences between speclmens assrgned 5v obliquely directed forwards (early meraspids) or s,tb- Rasetti to C. sardnusor C. ktus (see Rasetti, i972, p[. parallel for a short distance. Stout, bachvards direct- 18, figr 18-20, 25) and the specimens assignedto C. ed, genalspines, separated by ^more or lessimpressed delagei. Similar considerations about flattenirg, inter- furrow from the posterior margin of the fixigenae. nalGxternal mould morphology and paucir/ of the Librigenae unknowr, but probably tiny and narrow. available material are valid for the Tirrkish material (Shergold S( Sdzuy, 1984, p. 73-75, pl. 2, figr 9-1 1). Discussion In agreement with Geyer, earlv In agreement with Alvaro 6( Vizcaino (2000), corynexochids reveal a pronounced paedomorphic which discussed the position of the Sardinian til(a trend against dorypygids; these peculiar basal Mid- and prefer to maintain the species C. latus, we addi- Cambrian stock is in addition is distinct by a propar- tionally propose to restrict the name Corynexocltus ian suture, short palpebral lobes and a club'shaped latus to the rype material and assign the remaini.g more or less expanded glabella. The Sardinian Sardinian material to C. dekgei. For comparisons Clnuigellus is represented by only three meraspid with other described Corynexochusspecies ree R"setti cranidia,so, inappropriateto erect a new speciesand (1972), Courtessole (1973) and Shergold 6{ Sdzuy also to compare adequatelv with closer taxa. (1984, with references). However, it differs from the Moroccan C. annulu.rin 398 O. ELICI{I, G,L. HLLOIA having a lessvoulted cranidium, slenderglabella and Echinoderm plates,trilobites, brachiopods,bradori- more.-developedgenal spines, and alsd immature ids and hyoliihelminthids are exffemelyrare. So, fil- specimens a[*"yt" have ionger palpebral lobes; e ter feedeisare the clearly prominent group of the -.t"rpid of Chuigellusannulut (G.y.t, 1994, figs 7- deposidonal envirott-.ti it the begiini"L of tll. 17) clEarlyshows i slenderglabella, but fixigenaeare Campo PisanoFormation. For theseJreaniJms a rel- much more elevated. ativefyshallow and most possiblymudd| habitatwith A closely related taxon is represente.dby " -oderate water agitatiot ."t'be assumed(shallow Corynexochella?aenusta Dean (in Dean 6c Ozgi.il, subddal,e.g. MostlEr,1985). The depth was maybe 1994) from the basd Cal T.p. Formation (Thurus up to few tEnsof metres,what can be estimateddue Mountains, Turkey), which occurs at the baseof the t6 the intertidal to very shallow subtidal conditions Acadnparadoxida mureroensisZone, associatedwith during deposition of the directly underlayingstrata Protolenus (Pro to lenus) p isidianus . The Tirrkish species and to' thri ecologicalrequir.-.ttts of fauna ll There is clearly more similaf to the Moroccan and to the are no sisnsof allochthonousorigin for the spiculite Siberiantaxa than the Sardinianone. facies(as"'erosive base/planes, gttf,ittg, clasts).n fur- Ckaigelhu?n. sp. showsclose similarities with the ther argument for the lutochthonist. it suggesr..dby unassignEdCorynixochid specimen (Pl. 3, fig. 38, investigations-show on other Campo Pisano sections, from fevelGPI i); this latter is in turn closelyrelated which that this formation mostly starts with to the specimenassigned rc Corynexocltuscf. dekgeibv such a poriferid-rich portion. Basedon thesedata, it Sdzuy\gel, pl. 2i, fig. 7) *tr, as noricedby Geyer can surily be assumedthat fauna I is autochthonous. (1994), ffinI represent e proparian corynexochid The mendoned biotic monotony may reflect a cer- til(on close to Ckuigellus, bearitg ,,strettonia-like' tain kind of stress,most possibly ristricted water pdpebrallobes, present in the IberianChains (Spain), exchanseand/or oliqotrophic conditions. Montagne Noire (France)and now in Sardinia. The"following fain ^ i showsa distinct increasein echinoderms"tdtrilobite remains(mosdy agnosdds). Occurrence- Ckuigellus?n. sp. is restrictedto the Besides brachiopods,'The also chancelloriids and level GP2 of Gumur,I Pda sect:ion,which strongly hvolithids occur. latter are rather small and suggestcorrelation with the Protolenus(Protolenus) iriclude as well specimenswith circular cross-section, pili&ianus beds of the CPI assemblage(strata with- which areinterpr?ted a^s semi-infaund mud-stickersas quadrangular but Paradoxides)indicative of uppermost Early thosewith a flattenedbase or with cross- (com- Cambrian age. secdon,indicating an epibenthicmode of"tif. pare Elicki, 1994\. Chahcelloriids,qpical of shallow io middle-subtidal conditions, are representedby * PAIAEOECOLOGICALDISCUSS ION "ma>rimum" only low arnount with a little at the base of fauna,2. Most of the abovementioned fossil groups As in younger systems,also for the Cambrian a of fauna 2 have realiseda suspension-feedinebehav- compl.* of cott?itionscan be assumedwhich signifi- iour on the sedimentsoft- to firqground t,tt6.. Ora- cantly effectedthe biodc diversiryof assemblagesand chiopods: filter-feeders,agnosdds are mosdy repre- their'regional and ecologicaldiJtribudon. Alihough r.ntid by benthic deritus-salnplingsuspension- ind the pr.ft*ation potentill of organisms(and so th. deposit-feedersICondylopyge, Peronopsis and most pos- fossil record, tespectively)is different dependingon 'Waiossek,siblv also Dawsonial; fdi discussionsee Mtillei 6( the occupiedrypes ^Cambrianof habitats, it is wid ely accepted 1987; Debrenne 6c Zhuravlev, 1997; that during thi benthic assemblagesharre \il{hittington et al., 1997; Huehes,2001; Geyer,pers. dominated (Debrenne&. Zhuravlev, 1997). comm.)."In comparisonto faine l, this fauna2 indi- Genemlly, there are biodc and abiotic factors catesan increaseof suspensioninput into the open- which influencethe ecologicwindow. The main abi- marine shallowto middle-subtidal'conditions. otic factors for an environment were representedby The succeedingfauna 3 contains the samefossil nutrients, temperature, gradual l1ght,.o)rygenic level, available sroups as fauna I, but with a vanishing of chemistry of water and substrate,water energy and Eh"ttt.[oriids and hyolithids.TKe content on echin- clasticinput. Especi"lly,the three first mentionedare oderms and trilobites now decreasessignificandy. stronelv"'fr'"-.'ofi-portant "td setabsolute limits for the eco- Nevertheless,these two epibenthicgroups are the sdll logic"E*..pt an assemblage. dominating biota here.Vhereas the deposit-and sus- for one record 6f Ciruanelk published by pension-feEdirgtrilobites representiutochthonous Cherchf 6c Schroeder (1984), prim ^ry producers il.-ettts by t"ihot omic arguments,for the remains (bacteria,cyanobacteria, algae) are not known from of the t,rtp.nsion-feeding..f,ittoderms (distinctlyless the C"rnpo PisanoFormation until now.This is most than in faune 2) a move[rent from shallow and more (as in degosits, likely a taphonomic phenomenon, already known aqitated areasis possible - , young:r froni -"tty other carbonatedeposits of all epo_chs. where transportation of skeletal remains of this At Guffuru Pala,the rather monotonous fauna I organismgroup without markableagitation is possi- consists nearly exclusively of poriferid remains. bl; over ri-. ditt"ttce). This fauna-3persists over a C,UIBRAN MICROFAUNAAND PAI}IEOECOLOGYOF THE CAMPOPISANO... 399 large portion of the section and represents a stable ACKNO\TLEDGEMENTS open-manne, probably deep subtidai environment. ^ very for the SEM work to Claudio The following r,r.t.tsion of fauna 4 | fauna 5 | The authors are srateful Gendlini (Modena and'deggio Emilia Universiry) ald to Anja fauna 6 shows a quite similar pattern: first, passivefil- Obst (GeologicalInstitute, FreibergUniversiry). Further thanks ter-feeders predominate, followed by trilobite-echin- qoes to Rodolfo Gozalo (Valencii) for useful suggesdonson oderm facies and finally a decrease'of trilobites and eondllopygediscussion, to Gerd Geyer ()fli.irzbutg)-Tot remarks (I-i-lle) echinoderms (dominance of most possible deposit- otr "{.orila plaeoecoloBy and to JavierAlrraro and a fur- ther lttony^bus refere.ftrt helpful-remarks.The researchwas sig- feeders). The fossils of the last decimetres of the nificantly'supported by the G.t-"tt ResearchFoundation- (f6r "Comparative Campo Pisano Formation are rather small and fre- O.E.r t.i."r.h proiect gl- t 44lS srudieson facies final stageof a-Cambrianpladorm evo- quently- broken. and palaeoecol6gy'ofthe So, it seemsthat the successionof habitats docu- ludon: the smail'shellyfauna of"the Campo PisanbFm. / S\f- Sardinia")and by MIUR (G.L.P.). mented at Gutturu Pala were characterised twice by similar patterns: ( I ) firstly shallow subtidal, low-agi- REFERENCES tated and probably ecologically stressed(low diversi- followed by a tl"llo* to middle-subtidal, ry), .'(2) Arvnno, J.J.&.YtzcAINo, D., 2000, Nouvel assemblagedes trilo- open-marine envrronment which changgd fiqally (3) bites dans le Cambrien moyen de la nappe de Pardailhan into the more distal and quiet deeper subtidal. (Montagne Noire, France):implications biostratigraphiques dans la rdgion mdditerranienne: Eclogae Geologicae The dramatic facies replacement duritg the ffan- Helvetiae, 93: 277 -289. sition from the underlying San Giovanni Formation ANGELIN,N.P., 1854, PalaeontologiaScandinavica, Pars l, (Gonnesa Group) to the Campo Pisano Formation CrustaceaFormationis Tiansitionis, fasc. 2: Palaeontologia (IglesiasGroup) i"$icates a reorganisation of the sed- Scandinavica:2l-92. may imentary and ecological environment, which BnnneNDE,J., 1846, Notice prdliminaire sur le systbmeSilurien best explained by teclonic instabiliry of the platform et les trilobites de Boh€me.Hirschfeld, Leipzig: l-96. (subsidenceimpulse , tensional tectonics), most possi- BEcHsrAor, T. 6c BoNI, M., 1994, Sedimentological,strati- bly accompanied by " moderate relative sea-levelrise graphicaland ore depositsfield guide of the autochthonous (e.g. Bechstldt 6( Boni, 1994; Loi et dl., 1995; Cambro-Ordovician of southwesternSardinia: Memorie descrittivedella cartageologica d'Italia, vol. XLUII. Servizio Peiej6n et a1.,2000; Elicki, 2001). A climatic change GeologicoNazionale: 434 pp. in that time interval is under discussion. Based on BrNcrsoN, S., 1990,Hyolithelminth. In Benqtson,S., Conwa)' sedimentological, isotopic investiga- jliagen.1!1 1r1.d Morris, S., Cooper,B.J., Jell, P.A.& Runnegar,B.N. (eds), don, Cocozza &, Gandin (1990) assumea transition Early Cambrian fossilsfrom South Australia:Association o[ from arid (Santa Barbara Formation, early Gonnesa AustralasianPalaeontologists, Memoir 9: 186-190. Group) to tropical-humid (San Giovanni Formation, - 6aMIssnnzHEVSKI, V.V., 1981,Coeloscleritophora - a major upp.ei Gonnesa Group) *fhiT." generally,tidal-flat group of enigmatic Cambrian metazoans.In Taylor, M.E. (.d.)l environment. In contrast, Perej6n et al. (2000) agree Short p"apersfor the secondlnternational Symposium on the Cambrian System:U.S. GeologicalSurvey Open-File in the arid conditions at early Gonnesa time, but they Report,8l-743: 19-21. (but not exclude generally) a transition to trop- doubt BoNt, M. & BnrnssoNE,G., 1994,Su Zurfuru and Gutturu Pala ical-humid becauseof the strong diagenetic overprint Mines (SanGiovanni Formation) . InBechstldt,T. 6{ Boni, M. of the upper Gonnesa Group. Consequently, the cli- (eds),Sedimentological, stratigraphical and ore depos_itsfiefd matic situation during deposition of the Campo guide of the autoihthonous Cambro-Ordovician of south- Pisano Formation may be concluded as still arid or *estern Sardinia: Memorie descrittive della carta geologica d'Italia,vol. XLVIII. ServizioGeologico Nazionalez 349-355. tropical-humid. Regarditg the higher terrigenous 1886,Die Versteinerungendes Cambrischen content of the Campo Pisano Formation an (l) BonNrueNN, J.C., Schichtensystemsder Insel Sardinien nebst vergleichenden increase of fluvial transported material and, conse- Untersuchungen i.iber analoge Vorkommnisse aus andern quend% of a wetter climate, or (2) a movement of the Lendern. Erste Abteilung: Nova Acta der Kaiserlichen deposiiional areainto other palaeoclimatic (= palaeo- Leopoldinisch-CarolinischenDeutsche n Akademie der -149. geographic) regions or (3) both, is most likely. Naturforscher,51: I The intereiting phenomenon at Gutturu Pala, -, l89l , Die Versteinerungen des Cambrischen nebst vergleichenden that the ecological successionof faunas I l213 more or Schichtensystemsder Insel Sardinien lJntersuchungentiber analogeVorkommnisse aus andern (faunas lessfollows " i.otd time 41516)higher in the Lendern. Zweite Abteilung: Nova Acta der Kaiserlichen section, can be best interpreted as a moderate and Leopoldinisch-CarolinischenDeutschen Akademie der tempo rarv fall of the relative sea-levelwithin a short Naturforscher,56: 425-529. time. AftLr that sudden fall, the more or less same BurrEnnELD,N.J. & NIcHOLAS,C.J. , 1996.Burgess Shale T'pe ecological successionof an again subsiditg environ- preservationo[ both non-mineralizingand."shellv" Cambrian ment, respectively of a risen-relative sea-lEvel(from organismsfrom the MackenzieMountains. 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