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Italy): a Discussion on Its Morphology and Possible Mode of Life Autecology of selected fossi1 organisms: Achievements and problems. A. Cherchi (ed.). Boll. Soc. Paleont. Ital., Spec. Vol. 3, Mucchi, Modena, 1996, 145-158, 3 pls., 6 text-figs. The trilobite Giordunellu Bornemann, 1891 from Lower Cambrian of Sat-dinia, (Italy): a discussion on its morphology and possible mode of life Gian Luigi PILLOLA Dipartimento di Scienze della Terra Università di Cagliari KEY WORDS - Trilobita, Functional morphology, Palaeoecology, Twace ridges, Pits, Lower Cambnan, Sardinia, Italy ABSTRACT - The trilobite Giordanella Bornemann, 1891 (Dolero1enidae)fiom the Lower Cambrian Ne& Group of SWSardinia, is represented by two species: G. rneneghinii (Bornemann, 1883) and G. vincii Pillola, 1991 bothlshowing the earliest known examples of illaenid- like morphology. 7%e possibtlity that they spent part of their life in infaunal stance is suggested by the comparison with other trilobites with similar adaptivejèatures, some of which are claimed to bave been recovered in l+ position. Giordanella vincii irobablv, revresents ' an active ebifaunalL, trilobtte., livine close to calcimicrolnal-archaeocvathanmounds ofthe middle and upper Sa Tuvara Member (Matoppa Formation). It occasiomlly rnay bave assumed- a bumastoid stance inclined at low angle. The abundance of Giordanella rneneghinii in higher energy environments seem to be related to its pre-adapted morphology; its burrowing attitude may bave been exploited for protection against water turbulence, but also may bave been used for food gathering. The terrace ridge position and geometry show that the ventral ones and those on the dorsal stde on the pygtdium of G. meneghinii could bave functioned as sediment gripers. Those closely associuted with pits on the cephalic border may bave had a predominantly current-monitoring junction.-. RLASSUNTO - [Il trilobite Giordanella Bornemann, 1891 (Cambriano inferiore, SW Sardegna, Italia): considerazioni sulla sua mor- fologia e modo di vita] - Giordanella Bornemann, 1891 (Trilobita, Dolerolenidue), proveniente dal Gruppo di Ne& del Cambriano inferiore del SW della Sardegna, è il più antico esempio di trilobite illaeniforme finora noto; esso è rappresentato da due specie: G. meneghinii (Borne- mann, 1883) e G. vincii Pillola, 1991. La possibilità che questi due taxa trascorresseroparte della loro vita in posizione infaunale è suggerita dal confronto con altri trilobiti aventi morfologia e strutture adattative simili; fra questi ultimi alcuni sono stati ritrovati in posizione di vita. Giordanella vincii era probabilmente un trilobtte epibentonico, vivente in prossimità dei mounds a calcimicrobt e archeociatidi del Mem- bro di Sa Tuvara (Formazione di Matoppa), che occasiomlmente poteva assumere la posizione 'bumastoide' inclinata a basso angolo. L'abbondanza di G. meneghinii in ambienti a più elevata energia sembra legata alla morfologia preadattata; l'attitudine endobentonica parziale poteva essere sfruttata per una protezione rispetto alla turbolenza dell'acqua ed, eventualmente, per la ricerca di nutrimento. La posizione e la geometria delle terrazze cuticolari suggerisce che quelle situate sulla superjìcie esterna della duplicatura e quelle nei bordi della faccia dorsale del pigidio potrebbero aver avuto una funzione meccanica di tipo frizionale o stabtlizzante. Le twazze cuticolari intima- mente associate alle perforazioni localizzate al margine del ctphalon potrebbero aver espletato una funzione sensoriale predominante. INTRODUCTION formation and extends in the younger Monte Azzieddas Mb. and part of the Punta su Pranu Mb. Giordanella Bornemann, 1891, is a redlichioid of the Punta Manna Formation (Text-fig. 1). trilobite from the Lower Cambrian Nebida Group The diagnostic characters allowing distinction of the Iglesiente area (SW Sardinia, Italy). Th'1s genus between G. meneghinii and G. vincii are mostly is placed in a distinct subfamily (Giordanellinae located in the cephalic region: G. menegbinii has Pillola, 1991) belonging to the Dolerolenidae shorter palpebral lobes and a much longer distance Kobayashi 1951; this assignment is mostly based on (exsag.) between the posterior tip of the palpebral the strong similarity between the ontogenic develop- lobes and the posterior margin of the fixed cheeks. ment of Giordanella and Dolerolenus Leanza, 1949 This trend is more pronounced in larger cranidia (Pillola, 1991). and proves an allometric growth for the cephalic Two species of Giordanella have been retained parts mentioned above (Pillola, 1991, fig. 35, herein from the previous literature: G. menegbinii (Bor- Text-fig. 2). nemann, 1883), and relative synonyms (see Rasetti, Some morphological features seem to be peculiar 1972; Pillola, 1991), and G. vincii Pillola, 1991. to G. meneghinii: These two species do not coexist in any known sec- - occurrence of terrace ridges on the cranidial tion: G. vincii occurs in the middle and upper parts border, on the free cheek margins and on the latero- of the Sa Tuvara Member of the Matoppa For- dorsal regions of the pygidium (ventral terrace ridges mation, while G. meneghinii appears in the upper- on the doublure occur in both species). most beds of the Sa Pruixina Member of the same - occurrence of pits (= punctae, fine per- 146 G. L. PILLOLA Text-fig. 2 - Scatter diagram showing plot of the exsagittal length of the palpebra1 lobe (LLp) against the total sagittal cranidial length (LCr) in Giordanella meneghinii (Bornemann, 1883) and in G. vincii Pillola, 1991. Note: in the representation of the cranidium of G. vincii and of the cephalon of G. meneghinii the vertical mid-lines separates the in- ternal mould (left side) from the external surface (right side). After Pillola, 1991, modified. forations of the test), covering the whole dorsal carapace. The genera1 aspect of Giordanella and the typical features of G. meneghinii will be described below and discussed in comparison with other effaced trilobites, in order to propose a possible mode of life. The discussion of (1) the functional morphology (considering the balance between well adapted and - unfavorable features for a specific function) and also (2) the analogies with living arthropods and (3) the independent geologica1 information (see For- tey, 1985), may provide a reasonable and safer basis . for the deduction of the probable function of some features and, as far as ~ossible,the trilobite ecolog~. SEDIMENTARY AND FAUNAL CONTEXT The Cambrian sequence from SW Sardinia is composed of three main lithological units which are, in ascending order: - the Nebida Group (Atdabanian to Botomian), including mainly terrigenous clastics with microbial- archaeocyathan buildups (Matoppa Formation) and mixed siliciclastic and carbonate de~osits(Punta Text-fig. 1 - Schematic column of the Lower Cambrian Nebida Manna Formation); Group from eastern Iglesiente (SW Sardinia, ItaIy), - the Gonnesa Group (Botomian to Toyonian), showing the range of Gi~~~nel~avinci2 and G. consisting exclusively of carbonate sediments, with meneghinii. a) Sa Tuvara Mb.; b) Sa Pruixina Mb.; C) Monte Azzieddas Mb.; d) Punta Su Pranu Mb.; e) dolostones at the base (Santa Barbara Formation) , cuce"Aspu Mb.; 0 Biasterria ~b.~f~~~ and limestones (more or less dolomitized) at the top Pillola, 1991, rnodified. (San Giovanni Formation); THE TRILOBITE GIORDANELLA BORNEMA NN 147 - the lower part of the Iglesias Group (upper- Trilobites and other fauna1 remains occur more most Toyonian to Tremadoc), which begins with or less close to the carbonate mound systems. The the argillaceous or nodular limestones of the Campo trilobite communities are dominated by Meta- Pisano Formation (Toyonian-mid Middle Cam- redlichiinae (Iglesiella, Sardoredlicbia, Metaredlicbia) brian); this latter is overlain by the Cabitza For- and/or Dolerolenidae (Dolerolenus, Gior'danella) mation, represented by siltstones, sandstones and while Pararedlichiinae, Metadoxididae and eodiscids shales, and sporadic limestones, yielding, in its mid- are minor components. Other skeletal remains are dle part, a Lower Tremadoc fossi1 assemblage represented by Archaeocyatha, both inarticulate and (Pillola, 1990, 1991). rare articulate brachiopods, molluscs (Stenotbecoides, Giordanella is the commonest trilobite of the Hyolitha) and Coeloscleritophorans (Cbancelloria, Nebida Group and is widely distributed thorough Ez;ffelia?). the Iglesiente and Sulcis (SW Sardinia). G. vincii oc- During deposition of the Sa Pruixina Member an cur in the middle and upper part of the Sa Tuvara important terrigenous episode, related to a sand shoal Member (lower Matoppa Formation), N2 (pars) environment, stopped definitively the development trilobite Zone. G. menegbinii appear at the top of Sa of the Epipbyton-Renalcis dominated mounds. Later Pruixina Member (upper Matoppa Formation) con- on, this shallowing upwards sequence evolved into comitant with the lowermost occurrences of subtidal to pure oolitic shoal and back shoal environ- Dolerolenus longioculatus Pillola, 1991 and the disap- ments of the basal Punta Manna Formation. pearance of D. aff. courtessolei (N2 to N3 transition) The Sa Pruixina Member is generally poorly and extends in the younger Monte Azzieddas and fossiliferous, except near the top, where the very Punta Su Pranu Member of the Punta Manna For- rich Giordanella menegbinii beds occur. mation (Text-fig. 1). The last occurrence of G. The Monte Azzieddas Member consists of a menegbinii took place in the upper part of the N3 cyclic succession of oolitic-oncolitic grainstones Zone, just below the massive occurrence of
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