Albian Rudist faunas from Southern Italy: Taxonomic, biostratigraphic and palaeobiogeographic aspects

I

AI,BIAN R U D I S T F A U N A S F R O M S O U T H E R N ITALY: TAXONOMIC, B I O S T R A T I G R A P H I C AND PAI,AEOBIOGEOGRAPHIC ASPECTS

JEAN-PIERRE M A S S E , MAaDA G A L L O M A R E S C A & ELENA L U P E R T O

SINNI

MASSE J.-P., GALLO MARESCA M. & LUPERTO SINNI E. 1998. Albian Rudist faunas from Southern Italy: taxonomic, biostratigraphic and palaeobiogeographic aspects. [Les faunes de rudistes d'Italie mdridionale: aspects taxonomiques, biostratigraphiques et pal6obiog6ographiques]. GEOBIOS, 31, 1: 47-59. Villeurbanne, le 28.02.1998. Manuscrit d6pos~ le 05.05.1996; accept~ d6finitivement le 03.09.1996. ABSTRACT - Albian Rudist faunas from southern Apennines (Abruzzi, Matese Mounts) and Apula (Murge and Gargano), hitherto mainly considered Cenomanian, are dated after micropaleontological assemblages. They consist of 12 taxa belonging to the Requieniidae, Monopleuridae, Polyconitidae, Radiolitidae and Caprotinidae (?). Our study, even preliminary, calls attention to the specificity of the Albian fauna relatively to the Aptian and Cenomanian ones. In this respect the importance of the genus Kugleria and of particular species of Polyconites, Horiopleura and Eoradiolites is pointed out. The anatomy of Himeraelites is distinct from those of the Caprinidae. Some elements of this fauna are cosmopolitic whereas some have an Arabo-african or even Apulian significance. KEYWORDS: RUDISTS, LOWER CRETACEOUS, BIOSTRATIGRAPHY, TAXONOMY, PALAEOBIOGEOGRAPHY, SOUTH ITALY. R]~SUMt~ - Les faunes de rudistes de l'Albien de l'Apennin mdridional (Abruzze, Monts du Matese) et d'Apulie (Murges, Gargano) jusqu'ici surtout consid6r~es comme c6nomaniennes, sont dat6es de l'Albien d'aprgs les assemblages micropal6ontologiques associgs. Sont recens~s 12 taxons appartenant aux familles suivantes: Requieniidae, Monopleuridae, Polyconitidae, Radiolitidae et Caprotinidae (?). Nos r~sultats pr61iminaires, mettent l'accent sur l'originalit4 de la faune albienne vis-g-vis des faunes aptiennes et c6nomaniennes qui l'encadrent. De ce point de rue, l'importance du genre Kugleria et de certaines esp~ces de Polyconites, Horiopleura et Eoradiolites est soulign6e. Uanatomie de Himeraelites est diff6rente de celle des Caprinidae. Certains 61dments de la faune sont cosmopolites, d'autres ont une signification arabo-africaine ou plus strictement apulienne. MOTS-CLES: RUDISTES, CRI~TACI~ INFt~RIEUR, BIOSTRATIGRAPH1E, TAXONOMIE, PALt~OBIOGt~OGRAPHIE, ITALIE DU SUD.

INTRODUCTION Albian r u d i s t f a u n a s from S o u t h e r n I t a l y h a d b e g a n to be m e n t i o n n e d since t h e n i n e t e e n t h cent u r y in p a r t i c u l a r by Di S t e f a n o (1888, 1898) t h e n r e v i e w e d or c o m p l e m e n t e d by S c h n a r r e n b e r g e r (1901) a n d P a r o n a (1909). Until r e c e n t l y the m a i n p a r t of t h e s e f a u n a s w e r e a s c r i b e d to t h e C e n o m a n i a n . P o t e n t i a l l y this s i t u a t i o n h a v e been g r a d u a l l y modified in the course of s u b s e q u e n t d e t a i l e d b i o s t r a t i g r a p h i c studies m a i n l y based on micropaleontological analysis (e.g. Chiocchini et al. 1994). Besides t h e basic works p e r f o r m e d at the t u r n i n g of t h e c e n t u r y we can notice some m o r e r e c e n t local c o n t r i b u t i o n s which dealt about Albian r u d i s t faunas, p r o v i d i n g i n t e r e s t i n g new data. A m o n g t h e s e are those from Torre (1964) concer-

n i n g t h e M u r g e Baresi, r e f e r i n g e x p l i c i t l y to Albian rudists as well as t h o s e from Mainelli (1975) focusing on some Albian t a x a f r o m t h e Abruzzi and t h e Matese. T h e m o s t r e c e n t contribution from one of us t e n d to focus on t h e radiolitids (Gallo M a r e s c a 1993, 1994) of t h e Murge. The p r e s e n t c o m m u n i c a t i o n deals with: a s t r a t i g r a p h i c discussion on t h e s t u d i e d localities in order to d o c u m e n t the p r o p o s e d Albian age, t h e r e a p p r a i s a l of s o m e c o n t r o v e r s i a l t a x a concerning t h e i r i n t e r n a l o r g a n i z a t i o n a n d system a t i c position, new d a t a on poorly defined or poorly k n o w n forms. The foregoing is based on r e c e n t investigations: field studies, s a m p l i n g of rocks a n d fossils and

48 FIGURE 1 - Geographic sketch map of the studied regions in Southern Italy and the investigated localities, a: Abruzze, b: Matese, c: Gargano, d: Murge Baresi. Situation gdographique des rdgions prospectdes de l'Italie du S u d et des localitds dtudides, a: Abruzze, b: Matese, c: Gargano, d: Murge Baresi.

1

.

I

~Camp'telMat l° ese ~

micropalaeontologic analysis performed on Albian platform carbonates from Apula (Murge and Gargano) and the southern Apennine (Abruzzi and Matese Mounts) (Fig. 1).

GEOGRAPHIC AND STRATIGRAPHIC SETTING

~

%

6Km

M ~ Angelc

\

ir't° ?'~

~

~

well known Monte la Costa, extensively explored by Mainelli (1983), focusing on the Himeraelites bearing beds which provide Orbitolina (Mesorbitolina) sp. and Triploporella cf. marsicana PRATURLON. They overlie Offueria bearing beds (Masse 1992) of Lower Aptian age capped by radiolitid beds homologs to the Sbregavitelli ones (Masse et al. 1993) ascribed to the Late Aptian. GARGANO (Fig. lc)

ABRUZZI (Fig. la) The studied material was collected at the Colle Pagliare (Monti d'Ocre) near l'Aquila and corresponds to the "Pagliare fauna" described by Schnarrenberger (1901) or the "Calcari a Nerinea forojuliensis" described by Parona (1909). These f a u n a s were m a i n l y considered Albian by Schnarrenberger and Cenomanian by Parona. The assemblage closely ressembles those described by Di Stefano (1888, 1898) from Termini Imerese (Sicily) also considered Cenomanian. This assemblage was found in the "Calcari a Polyconites verneuili and the Calcari a Caprotina" now regarded mainly Albian (Camoin 1982) after the orbitolinid association and because the corresponding beds are bracketed by well dated Aptian and Cenomanian sequences. Among the rich assemblages described by our predecessors we will concentrate on Himeraelites, complementary data will be also given on some associated forms of Requieniidae, Polyconitidae and Radiolitidae. MATESE (Fig. lb) Besides the former investigated localities (Masse et al., 1993) of Aptian age, we have studied the

Our material derives from two localities. First the breccia beds from the upper part of the M a t t i n a t a limestones Formation, of Late Albian age (Luperto Sinni & Masse 1987). The collected samples are from the marly matrix surrounding blocks as well as from some blocks themselves, thought to be of similar age because they yield a similar fauna. S o u t h w a r d to the Gargano massif, Albian sequences have been investigated u n d e r n e a t h the bauxite horizons at the locality casa Lauriola. The substratum of the bauxite is assigned to the Lower Cenomanian with alveolinids (Sellialveolina vialli COLALONGO and Ovalveolina maccagnoae DE CASTRO), the underlying limestones consist of requienid/monopleurid beds with some radiolitids, associated with algalaminated wackestones. These beds contain: Nummoloculina sp., Praechrysalidina infracretacea LUPERTO-SINNI(advanced forms with a cribate aperture), Cuneolina pavonia D'ORB., Mayncina bulgarica LAUG & PEYBERNES, Sabaudia minuta (HoFKER) and various representatives of Rumanoloculina. Of special interest is the presence of Cribellopsis arnaudae CHIOCCHINI and Neoiraqia cf. insolita

49 (DECROUEZ •

RV

MOULLADE) which m a r k the Albian

(Chiocchini et al. 1994). MURGE BARESI (Fig. ld) J

The studied localities pertain to the so-called "Livello Palese" (sensu Valduga 1965) outcropping on the coastal region between Bari and Giovinazzo and classically regarded as Albian (Luperto-Sinni 1966). This region also includes the rudist bearing beds studied by Torre (1965) at Bitonto and Molfetta farther from the coast. The age of the Livello Palese in these localities is tentatively regarded to be Middle to Upper Albian. This assignment is based on the presence of Neoiraqia cf. insolita (GALLOMARESCA,1994) (Fig. 4.1,2).

SYSTEMATIC PALAEONTOLOGY Family REQUIENIIDAE Douvill6, 1914 Genus R e q u i e n i a MATHERON,1842

Requienia migliorini TAVANI,1948 Fig. 4.3.

The attributes of this form and its synonymy have been discussed earlier (Swinburne & Masse 1995). The main question regards the characters of the left valve: poorly known in the type specimens from Tavani (1948) and mainly documented from later synonyms: Requienia tortilis MAINELLI and Requienia polygyra (ALENCASTER).As noted by Chikhi Aouimeur (1980) the forms recovered from Mexico and Algeria are near identical whereas the forms from Italy have a more elongated, low assymetric conical shape. This taxa have been recorded from Casa Lauriola. It is also well represented in Late Aptian-Albian p. p. sequences from the Lago del Matese.

1 cm i

FIGURE 2 - Toucasia sp. (Casa Lauriola, Gargano). Dorso-ventral section of a bivalve specimen showing the main myocardinal elements: pmp- posterior myophoral plate, pt- posterior tooth, ps- posterior socket (RV- right valve). Section dorso-ventrale d'un spdcimen bivalve montrant les principaux dldments myocardinaux: prop- lame myophore postdrieure, pt- dent post~rieure, ps- fossette postdrieure (RV- valve droite).

(illustrated Fig. 4.a-d) the posterior side of the right valve is convex outward and seems to project outside the commissural plan, as in Toucasia transversa. We consider the small size (the umbo/ventral side length fluctuates from 3 to 4 cm) a significant attribute of this taxa (whereas Toucasia transversa is commonly larger t h a n 8-10 cm). The myophoral plates of Toucasia transversa tend to parallel the commissure and intersect on the postero-ventral part of the shell. On the specimens from Torre and on our own material the intersection of the myophoral lines on the commissure are located in the middle part of the shell and the myophores intersect with a higher angle. Moreover, the ventro-dorsal section of bivalves specimens show, as in Toucasia transversa: - a relatively small posterior socket on the left valve corresponding to an erect near vertical posterior tooth of the opposite valve (Fig. 2), - a wide extent of the cardinal platform ventrally, giving to the internal molds a comarginal bulge lining the commissure (Fig. 4.4). This form could be regarded as a new species. Genus Kugleria BOUWMAN

Genus Touvasia MUNIER-CHALMAS,1873

Kugleria steinmanni (SCHNARRENBERGER, 1901) Fig. 3, 4.5,6

Toucasia sp. Fig. 2, 4.4

The genus Toucasia is represented by small size species in various localities (Murge Baresi, Casa Lauriola). This material is partly equivalent to those described as Toucasia transversa var. minuta by Torre (1965). The authors' opinion is t h a t the fauna described by Torre correspond to at least two distinctive taxa. The first form (illustrated on Fig. 4.4a-b) is close to Toucasia compressa MASSE (Masse 1976) t h a t is to say r a t h e r distinct from Toucasia transversa PAQUIER. On the second form

This

taxa

formerly

described

as

Toucasia

(ScHNARRENBERGER, 1901) was transfered to the

genus Kugleria proposed by Bouwman (1938). As noticed by this author the genus is strongly homeomorph with Toucasia but the posterior myophoral plate at the right valve is just an extension of the cardinal platform and not an independant lamina as in Toucasia, whereas there is a posterior myophoral plate on the left valve. Moreover the anterior myophores display a high convexity and the posterior tooth projects

50

RV

/ cp / pmp

1 cm

FIGURE 3 - Kugleria steinmanni (Colle Pagliare, Abruzzi). Antero-posterior section of the right valve showing the posterior myophoral plate (pmp) representing the extent of the cardinal platform (cp) and the anterior myophoral bulge (amb). Section antdro-postdrieure de la valve droite montrant la lame myophore postdrieure (prop) reprdsentant l'extension du plateau cardinal (cp), et le bourrelet myophore antdrieur (arab).

obliquely outside the shell. The Italian species departs from the Caribbean one (K. macgillarryi BOUWM~N) by the acute posterior margin of the right valve. Our material derives from the Colle Pagliare that is to say the locality where this taxa was formerly mentionned by Parona (1909). The so called Matheronia? ausonicola described by Parona from the same locality could be also a representative of the genus Kugleria because all the attributes of this taxon are found in the former

one.

Family MONOPLEURIDAE Munier-Chalmas This family is represented by tubular shelled forms tentatively ascribed to Petalodontia POCTA. Because our material is exclusively composed of right valves a generic assignment remains problematic. At least two forms could be distinguished. The first one displays fine concave upward tabu-

lae in the body cavity (Fig. 4.7). The second one lacks both the external costulation and the tabulae (Fig. 4.8). These forms closely resemble Petalodontia sp. 2 and Petalodontia sp. 1 respectively described by Chikhi Aouimeur (1980) from the Late Aptian of NE Algeria. Forms belonging to the same group are also found in the Late Aptian of the Matese Mounts (personal observations of JPM). The presence of Agriopleura? darderi ASTRE (a probable synonym from Mathesia tertiicolloquiirudistarum MAINELLI) cannot be ascertained. Our material is densely bored and the typical tubular microstructure of the outer calcitic shell layer (see description in Masse & Philip 1974 and Chikhi Aouimeur 1980) deeply altered. The so called "calcari con anelli" (= limestones with rings), reported from various author in southern Italy, does not only correspond to "Mathesia tertiicolloquirudistarurn" as suggested by Mainelli (1995) but also to the "Petalodontia" group. Family CAPROTINIDAE?Deschaseaux p.p. Genus H i m e r a e l i t e s DI STEFANO, 1888 Fig. 4.10,11

This genus is a major contributor to the Albian rudist fauna in the Monte d'Ocre (Parona 1909) and at Monte la Costa. Because its anatomy and systematic position have been subjected to contrasting interpretations we intend to reappraise the myocardinal organization: a basic approach to relate this genus to the close comparable "Caprotina"-Sellaea group of Di Stefano (18881898). Himeraelites was first proposed by Di Stefano (1888) as a s u b g e n u s of t h e genus Monopleura MATHERGN.Later on it was regarded

FIGURE 4 - 1-2. Neoiraqia cf. insolita Giovinazzo (Murge Baresi). 1, axial section [x 30]. 2. Transverse oblique section [x 25]. 3. Requienia migliorinii (Abruzzi). Left valve showing the helieospiral coiling. 4. Toueasia sp., Casa Lauriola (Gargano). Posterior view of the internal mold showing the tracks of the myophoral plates and the cardinal platform (underlined). 5,6. Kugleria steinmanni (Colle Pagliare, Abruzzi). 5, view of the right valve showing the coiling habit. 6. posterior view of the same valve showing the absence of any external myophoral groove: the posterior tooth (pt) is broken. 7,8,9. Petalodontia? sp. Casa Lauriola (Gargano). 7-8, tabulate form. 7, longitudinal section showing the concave tabulae (t). 8, transverse section. 9. non tabulate form, transverse section: the inner, initially aragonitic, shell layer have been dissolved and the corresponding void filled with a muddy sediment (arrow). 10,11. Himeraelites sp. (Colle Pagliare, Abruzzi). 10. internal view of the left valve showing the main myocardinal elements: at- anterior tooth, pt- posterior tooth, cs- central socket, am- anterior myophore, bc- body cavity; the posterior myophore (pm) lies on the ventral inner portion of the central socket. 11. ventral view of the right valve showing the central tooth (ct) and the adjacent bulge representing the posterior myophore (pro). 12. Eoradiolites plicatus (Colle Pagliare, Abruzzi). Transverse section of the right valve showing the ventral elements: ventral rib (v), anterior (AB) and posterior bands (PB). The interpretation of this section is also given on text-figure 8. Scale bar: 1 cm. 1. Section axiale 2. Section transversale oblique. 3. Valve gauche montrant l'enroulement hdlicospirald. 4. Vue postgrieure du moule interne montrant la trace des lames myophores et d u plateau cardinal (soulignd). 5. Vue de la valve droite montrant la disposition spiral@. 6. Vue postgrieure de la re@me valve montrant l'absence de sillon myophoral externe, la dent postdrieure (pt) est brisde. Z Section longitudinale montrant les tabulae concaves (t). 8. Section transversale. 9. Section transversale d'une forme non tabulde; la couche interne du test, initialement aragonitique, a dt@ dissoute, le vide correspondant combld p a r un sddiment boueux (fl@che). 10. Vue interne de la valve gauche montrant les principaux glgments myocardinaux: at- dent antdrieure, pt- dent postdrieure, cs- fossette centrale, am- myophore antdrieur, bc- cavitg gdndrale; le myophore postdrieur (pro) s'dtend sur le bord ventral interne de la fossette centrale. 11. Vue ventrale de la valve droite montrant la dent centrale (ct) et le bourrelet adjacent repr@entant le myophore postdrieur (pro). 12. Section transversale de la valve droite montrant les gldments ventraux: cr@te ventrale (v), bandes antgrieure (AB) et postdrieure (PB). Einterprdtation de cette section est reprise dans la figure 8. Echelle: 1 cm.

51 i~!i ~ii!~ i

52 an independant genus, the type species of which being Himeraelites vultur DI STEFANO(Di Stefano 1989). Himeraelites was considered to have a vertical posterior myophoral plate on the left valve and an horizontal anterior myophoral plate on the right valve, this valve also displays a posterior accessory cavity corresponding to the myophoral plate of the opposite valve. Douvill~ (1900) noticed that the muscle attachment onto the posterior myophoral plate was on its external side as in Petalodontia. Nevertheless in the Di Stefano's as well as in the Parona's interpretations the muscle attachment is regarded internal to this plate and located on its ventral termination inclined toward the central socket. This interpretation was also shared by MacGillavry (1937) who underlined the affinity of this myophoral organization with its "caprinid type". Actually in contrast with the Caprinidae, Himeraelites lacks an erect myophoral plate projecting from the right valve into a myophoral cavity of the opposite valve. The posterior myophore of the right valve is near flat in its posterior portion, grading to a bulge ventrally: this bulge takes place on the dorsal (i.e. inner) margin of the "myophoral plate" of the left valve. This so called "myophoral plate" is just the elevated, marginal portion of the cardinal platform and has no myophoral function in contrast to Douvill6 opinion. The so called "Caprotina" (e.g. Caprotina strix) figured by Di Stefano (1898) as well as Sellaea from the same author display a close related myophoral organization which departs from those of Caprotina D'ORBIGNYreported b y Douvill~ (1886). In "Caprotina" (sensu Di Stefano) and Sellaea the posterior myophore of the left valve protrudes deeply in a myophoral cavity of the right valve, a configuration close to those of the Coalcomaninae (Chartrousse in press).

Himeraelites can therefore be regarded the root of the "Caprotina"-Sellaea group. Its myophoral organisation departs from those of the Monopleuridae and the Caprinidae. It also differs from the organization of the Caprotinidae with which it shows the strongest affinity. Our material, mainly collected at the Colle Pagliare belongs to the group of Himeraelites transversus PARONA and Himeraelites meghistoconcha DI STEFANO.These forms are characterized by a wide subtriangular central socket and a relatively high antero-posterior development of the shell. These specific assignments are conservative, owing to the fact that the specific systematic framework of Himeraelites is in the author's opinion, unsatisfactory because based on a limited number of specimens precluding to evaluate the range of intraspecific variability.

am

1 ~m

as

V

pm

FIGURE 5 - Polyconites cf. foveolatus (Colle Pagliare, Abruzzi). Inner view of the right valve showing: the body cavity (BC), the anterior (am), and posterior myophores (pro), the anterior (as) and posterior sockets (ps), the ligamentary crest (LC), the Iigamentary groove (L) and the calcitie outer shell layer (cl). Vue interne de la valve droite montrant: la cavitd gdndrale (BC), le myophore antgrieur (am), le myophore postgrieur (pro), les lossettes antdrieure (as) et post~rieure (ps), l'ar~te ligarnentaire (LC), le sillon ligamentaire (L) et la couche calcitique externe (cl).

Family POLYCONITIDAEMac Gillavry, 1935 The attributes of the two genera of this family:

Horiopleura MUNmR-CHALMAS and Polyconites ROULLa~Dhave been discussed by Douvill~ (1889), Di Stefano (1889) and Mac Gillavry (1935). The presence of Polyconites was reported from the Colle Pagliare fauna by Parona (1909) who described three species tentatively ascribed to this genus. Di Stefano (1889) also mentioned several distinctive species of Polyconites in Sicily. Horiopleura was not reported in both regions. We have collected representatives of the Polyconitidae in four localities: Monte San Angelo, Monte La Costa, Giovinazzo and Colle Pagliare. From Monte La Costa and Monte San Angelo the sampled material does not allow us to separate precisely Horiopleura from Polyconites. Two forms, with distinctive characters will be described more precisely. Genus Polyconites ROULLAND, 1909

Polyconites cf. foveolatus PARONA Fig. 5

A single specimen of right valve was collected at Colle Pagliare, assigned to Polyconites? foveolatus described from the same locality by P a r o n a (1909). This species was mainly defined after a single left valve characterized by a subcircular outline, externally flat, two teeth with a well developed longitudinal costulation, a thin fiat anterior comarginal myophoral plate, with an indentation which separates the teeth from the myophore; the posterior myophoral plate is elliptical and orthogonal to the teeth axis. The right

53

valve of our collection shows an elliptic outline in transverse section, the commissural plan is deeply inclined dorsally to the main axis of the shell, the dorsal side possesses fine ribs, the ligament groove corresponds to a ligamentary crest, the two sockets are wide, the posterior displays internal ribs, the posterior myophore is narrow, depressed and deeply inclined inward, and the anterior myophore is elongated and comarginal. Consequently, owning to the good correspondence between our material and those described by Parona, we can ascribe our specimen to the above mentionned species. This assignment tends to confirm the generic name proposed by Parona. Genus Horiopleura MU-NIER-CHALMAS

Horiopleura aff. almerae

PAQUIER

pm-%

pmJ' BC~ a m cl~~ al

Fig. 6 1 cm

From Giovinazzo (I1 Crocefisso) have been encountered several specimens of Horiopleura evidenced after random or oriented sections. The diagnostic attributes of this genus have been recognized after longitudinal antero-posterior sections showing: the external rounded, convex outward left valve, with a pediculate posterior myophoral plate and a convex downward anterior myophore; - the near horizontal, concave upward, posterior myophore of the right valve, the anterior one is slightly inclined inward. -

Transverse sections show a subcircular outline and on the right valve the thick internal shell layer corresponding to the myophores.

FIGURE 6 -Horiopleara aff. almerae (Giovinazzo, Murge Baresi). Longitudinal, antero-posterior sections of a bivalve specimen showing on the left valve the pediculate posterior myophore (pmp) and the anterior myophoral thickening (am). The right valve shows the myophoral transverse thickenings, the posterior being depressed (pro). The calcitic outer shell layer (cl) is thin and the inner shell layer, initially aragonitic (aI), relatirely thick. Sections longitudinales, antgro-postdrieures d'un spdcimen bivalve montrant sur la valve gauche le myophore postdrieur pddiculd (pmp) et l'gpaississement myophoral antdrieur (am). La valve droite montre les dpaississements myophoraux transversaux, le postdrieur gtant dgprimd (pro). La couche calcitique externe du test (cl) est mince et la couche interne, initialement aragonitique (al), relativement dpaisse.

foveolatus: two features which have been observed in the material from Giovinazzo.)

The diameter fluctuates from 6 to near 10 cm. The absence of external ribs, radial bands and of an axial ridge ontop of the left valve associated with the convexity of this valve and the shape of the posterior myophore of the right valve suggest a close affinity with Horiopleura almerae PAQUIER (Paquier 1903). Polyconites distefanoi described by Parona (1909) from the Colle Pag]iare shows also some similarities. This species possesses the myophoral attributes of Horiopleura instead of Polyconites clearly expressed on the figures provided by Parona on plate XXI. A significant feature is illustrated on figure 8: a longitudinal oblique section showing the horizontal concave upward posterior myophore of the right valve while the transverse sections (Figures 8-9) display a thick inner shell layer (right valve) and a strong myophoral apparatus (left valve). This form possesses a pronounced radial, anterior depression on the left valve, corresponding to the "anterior band", moreover the connexion between the posterior myophora] plate of the left valve and the adjacent teeth is indented (as in Polyconites

Polyconites sp. Fig. 7

From the Horiopleura bearing beds of Giovinazzo, we have also observed a poorly preserved material tentatively assigned to Polyconites. This assignment is based on oriented sections showing: - on the left valve (Fig. 7) the anterior cone with a myophoral meaning, the central cone corresponding to the body cavity and two small cones corresponding to the posterior myophoral zone; - on commissural section of the right valve the two teeth of the opposite valve and a wide body cavity: a pattern which suggests a thin, weak myophoral apparatus. This organization matches those of Polyconites. Family RADIOLITIDAEGray, 1848 As reported earlier (Gallo Maresca 1993, 1994) this group is only represented by the genus

Eoradiolites.

54

V \

\

pml I

/11

"

am

t

cl

/ . \

1 cm

a

1 cm

FIGURE 7 -Polyconites sp. (Giovinazzo, Murge Baresi). Outside view of the internal mold of a left valve showing the tracks of the anterior myophoral plate (amp), the central lobe (c1) corresponding to the body cavity, the cones 1 and 2 related to the posterior myophoral lobe (pml), the calcitie outer shell layer (cl) is only partly preserved. Vue externe d'un moule interne de la valve gauche montrant la trace de la lame myophore antgrieure (amp), le lobe central (cl) correspondant h la cavit# gdndrale, les c6nes I e t 2 rattachds au lobe myophoral postdrieur (pml); la couche calcitique externe du test (cl) n'est que partiellement pr~servde.

Eoradiolites plicatus (CONRAD) Fig. 4.12, 7

This species is documented from a single transverse commissural section of the right valve found at Colle Pagliare in the Himeraelites beds. The attributes of this form include: the convex outward growth lines of the radial bands, this structure which matches with the external morphology is typical of the genus Eoradiolites as shown by Gallo Maresca and Masse (1993); a subrectangular transverse outline; the peculiar habit of the "cellular structure": essentially meandriform with a relatively small number of wide rows, this cellular structure is mainly restricted to the dorsal side and ponctually in the posterior ventral, radial band; the shape of the radial bands: strictly confined to the ventral side, with a near flat outer surface; -

/

V

I

AB

\PB

FIGURE 8 - Eoradiolites plicatus (Colle Pagliare, Abruzzi). Transverse section of the right valve showing the subquadrangular outline, the location of the cellular/meandriform shell structure. V- ventral rib, AB- anterior band, PB- posterior band, LC- ligamentary crest; the myocardinal elements of the left valve: am-anterior myophore, pro- posterior myophore, are in grey. Section transversale de la valve droite montrant l'allure subquadrangulaire et la localisation de la structure cellulaire/mdandriforme du test. V- cr~te ventrale, AB- bande antdrieure, PB- bande postdrieure, LC- ar~te ligamentaire; les dldments myocardinaux de la valve gauche: myophore antgrieur (am) et myophore postdrieur (pro) sont en gris.

the posterior is thicker than the anterior b u t narrower; the dimensions conform to those given by Gallo Maresca (1993) for the type material; the "ventral crest" (V) is well marked; the ligamentary crest is subrectangular/trapezoidal; the myocardinal apparatus (still preserved in situ) conforms to those of Eoradiolites with a horse-shoe aspect in transverse section.

-

-

Eoradiolites lyratus (CONRAD) Fig. 9.1-4

Parona (1909), Douvill6 (1910, 1913) and Gallo Maresca (1993, 1994) have pointed out the outer and inner attributes of this species.

FIGURE 9 - 1,4. Eoradiolites lyratus (Monte Sant'Angelo, Gargano). 1, longitudinal section of the right valve showing the structure of the calcitic shell layer, compact and/or cellular at different ontogenetic stages. 2. Microscopic aspect of the compact shell layer of the right valve displaying the growth banding, Ix 20]. 3. Posterior view of the right valve, showing the ventral rib (V), the anterior (AB) and the posterior (PB) bands. 4. Transverse section of the right valve; V- ventral rib, AB- anterior band, PB- posterior band, L- ligamentary crest [x 1]. Black scale: 1 cm. 5,7. Eoradiolites murgensis. 5. Lauriola (Gargano), microscopic aspect of the shell structure in the posterior band showing the interbedded compact and cellular habit, Ix 20]. 6. Giovinazzo (Murge), view of the posterior right valve of the ventral rib (V), the anterior (AB) and the posterior (PB) bands. 7. Giovinazzo (Murge), transverse section of the right valve. Same symbols as on fig. 7. Black scale: 1 cm. 1. Section longitudinale de la valve droite montrant la structure de la couche calcitique, compacte et / ou cellulaire & diffdrents stades de l'ontogen~se. 2. Aspect microscopique de la structure compacte du test de la valve droite montrant les bandes de croissance (lame mince). 3. Vue postdrieure de la valve droite montrant la cr~te ventrale (V), ainsi que les bandes antdrieure (AB) et postdrieure (PB). 4. Section transversale de la valve droite; V- cr~te ventrale, AB- bande antgrieure, PB- bande postdrieure, L- ar~te ligamentaire. Echelle noire: 1 cm. 5. Aspect microscopique de la structure du test au niveau de la bande postdrieure, montrant l'intercalation des couches compactes et cellulaires. 6. Vue postgrieure de la valve droite montrant: la cr~te ventrale (V) ainsi que les bandes antgrieure (AB) et postdrieure (PB). 7. Section transversale de la valve droite; m~mes symboles que sur la fig. 7. Echelle noire: 1 cm.

55

5

PB

56 Our specimens, collected at Monte Sant'Angelo (Gargano) from the breccia beds, show: - a cylindro-conical right valve with subcircularrectangular outline in transverse section; - an outer ornamentation consisting of numerous longitudinal ribs; - the ventral rib "V" weakly prominent; - the radial bands on the posterior side, with flat or slightly concave outer surface and, between them, a smooth narrow groove; the anterior band is wider t h a n the posterior but thinner; - a trapezoidal ligamentary crest (transverse section). The most characteristic feature of our specimens is the structure of the calcitic outer shell layer. In some specimens, the entire shell consists of a subquadrangular cellular type, sometimes interbedded with a compact structure; the latter is frequent in the early ontogenetic stages but it has also been found in the later ones. In some specimens, the compact structure is dominant while the cellular one m a y be absent or has only a limited thickness.

Eoradiolites rnurgensis TORRE, 1964 Fig. 9.5-7

From the Murge (near Palese and Bitonto) Torre (1964) documented the presence of Eoradiolites murgensis TORRE and Eoradiolites davidsoni (HILL) while Campobasso (1972) referred some specimens from Giovinazzo to Radiolites peroni (CHoFFAT). Gallo Maresca (1993, 1994), in a study on Albian Radiolitids from Murge and Gargano, ascribed the specimens of Eoradiolites davidsoni and Radiolites peroni to Eoradiolites murgensis. Several specimens of Eoradiolites have been collected at Giovinazzo (in the same locality as Campobasso 1972) and in spite of their bad state of preservation, it was possible to recognize the diagnostic attributes of Eoradiolites murgensis consisting of: a flat left valve with a low convexity in the central part; the conical right valve with numerous longitudinal ribs; a transverse section with a triangular to elliptic outline; the ventral rib "V" is well marked; - the groove between the ventral rib and the anterior band is 3-10 mm wide, 1-3 mm deep; - the radial bands are mainly developed on the ventro-posterior side, with morphometric relationships (anterior band wider than the posterior but less thick) typical of the genus Eoradiolites (Douvill6 1909; Gallo Maresca 1993); the groove between the bands is wide (about 5-10 mm), deep (about 2-7 mm) with one or two small adjacent ribs; -

- the ligamentary crest is very small, with a trapezoidal shape (transverse section); - the myocardinal apparatus conforms to those of

Eoradiolites; - the calcitic shell layer shows a subquadrangular cellular structure lined, outwards by a compact structure which can be sometimes interbedded in the cellular one. At Casa Lanriola, we collected a single specimen of the right valve of Eoradiolites whose morphometric features are very close to those of Eoradiolites murgensis, the only difference being in the structure of the calcitic shell layer t h a t is mainly compact with a meandriform cellular structure especially in the posterior band. In the authors' opinion the radiolitids reported from the Colle Pagliare by P a r o n a (1909), Radiolites macrodon (PIRONA) and Praeradiolites pironai PARONAare likely to be transfered to other distinctive genera. The figures provided for Praeradiolites pironai merely conform to Eoradiolites t h a n to Praeradiolites. The attribution to Radiolites of a single isolated left valve having a convex u p w a r d shape appears unreliable. Consequently these forms m u s t be carefully revised in the future.

DISCUSSION At family and generic levels the above mentionned assemblage closely resembles the Albian one reported from the South of France (Masse, 1995).

Horiopleura, Polyconites, Eoradiolites, Agriopleura? (= Mathesia) and Toucasia being typical members of the corresponding fauna. The absence of

Pseudotoucasia: a common form in the Late Aptian-Albian sequences of northwestern Europe (Masse 1995; Masse et al. in press) is remarkable. Conversely the presence of Himeraelites, a classical form of the Apulian and African domains (Masse & Philip 1986) is stricking. This r e m a r k also works for Kugleria which n e v e r t h e l e s s appears to be widely distributed, from the New World (Bouwman 1938) to the Pacific (Yabe Guyot, off Japan) (Shiba 1979). During the course of our investigations we did not record significant representative of"Caprotina" and Sellaea, expected to have a stratigraphic and biogeographic extent similar to Himeraelites. At specific levels the similarities are less pronounced. Actually, the Toucasia and some representatives of Polyconites or even Horiopleura look distinct. The closest analogy is for the Eoradiolites species with an exception: Eoradiolites murgensis which seems to be restricted to the Apulian domain (Gallo Maresca 1993, 1994).

57 Similarly Kugleria steinmanni seems to be confined to the same area. From a biostratigraphic point of view the above mentionned assemblage, mainly regarded Cenomartian by earlier workers, proves useful for the identification of the Albian. Nevertheless the precise vertical distribution of its constitutive elements within the Albian, is difficult to assess. Our investigations only dealt with vertically limited sections with a m i n i m u m micropalaeontological control. Moreover our faunal record in only partial and regarded therefore incomplete (see above the c o m m e n t s on "Caprotina" and SeUaea). Di Stefano's work at Termini Imerese (Sicily) shows the existence of well d i f f e r e n t i a t e d vertical assemblages regarded stratigraphically significant. These observations and an incomplete record of the Colle Pagliare fauna reported by Parona (1909) suggest the possibility to find stratigraphical distinct faunas within the Albian sequences. Considering the relatively long duration of the Albian 15 My (Obradovitch 1993) and the existence of well known faunal changes during this timespan, documented for both shallow water platform biotas, as the benthic foraminifera (Arnaud-Vanneau et al. 1985), American rudist faunas (Scott 1990) and the nektonic or planktonic biota the biozonation of which are the foundation of the Albian biochronological framework (Haq et al. 1988) similar changes in the Italian rudist faunas could be expected, with a Mediterranean potential.

Moreover by discussing the affinities of the later species with Polyconites distefanoi, we suggest to regard this form an Horiopleura r a t h e r t h a n a Polyconites. Three species of Radiolitidae: Eoradiolites murgensis, Eoradiolites lyratus and Eoradiolites plicatus are recorded and some of their anatomic attributes specified. We notice the absence of Eoradiolites davidsoni. From a palaeobiogeographical point of view some elements of this assemblage are cosmopolitic (e.g.

Requienia migliorinii, Agriopleura? darderi, Eoradiolites plicatus) others as Himeraelites have an Arabo-african meaning, while forms as Eoradiolites murgensis and Kugleria steinmanni could be used as regional Apulian markers. Our overview, even preliminary, calls attention to the specificity of the Albian faunas, mainly considered Cenomanian by earlier workers. It is import a n t to recall however t h a t detailed taxonomic studies are still needed concerning the Himeraelites - "Caprotina" and Sellaea group, as well as on some representatives of the Monopleuridae and Polyconitidae. From the search for distinctive stratigraphical assemblages could be also expected a vertical biozonation and a significant knowledge improvement of the evolutionary patterns having an i m p o r t a n t bearing on the origin of m a n y Cenomanian faunas. A c k n o w l e d g e m e n t s - We t h a n k Mrs. G. H a u f e u r t who typed the manuscript and J.J. Roccabianca who printed the photographs. We are indebted to G. Sirna and D. Sartorio for their critical review of the manuscript.

CONCLUSIONS Rudist bearing rocks of Albian age investigated in southern Apennines (Abruzzi, Matese Mounts) and Apula (Murge and Gargano) are dated after micropalaeontological assemblages, mainly orbitolinids belonging to Neoiraqia and Cribellopsis. The Requieniidae include Requienia migliorinii, Kugleria steinmanni and Toucasia (pro Toucasia transversa var. minuta, p. p.) t h a t could be ascribed to a new species. The Monopleuridae are poorly defined, and tentatively assigned to Petalodontia (?) sp. and Agriopleura ? darderi. Himeraelites hitherto considered a member of the Monopleuridae or the Caprinidae is regarded close to the Caprotinidae. This is based on the reinterpretation of the myocardinal organization, pointing out the absence of true erect myophoral plates on both valves. The Polyconitidae include Polyconites cf. foveolatus, the generic position of which is confirmed and Horiopleura aff. almerae. Horiopleura is therefore reported for the first time from southern Italy.

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