HARVARD UNIVERSITY
Ernst Mayr Library of thè Museum of Comparative Zoology
Qsc
MEMORIE
della Società Italiana di Scienze Naturali e del Museo Civico
Volume XXXII - Fascicolo I di Storia Naturale di Milano
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
UBR$« SEP »’
POLYCHAETE, SIPUNCULAN AND ENTEROPNEUST WORMS FROM THE LOWER CALLOVIAN (MIDDLE JURASSIC)
OF LA VOULTE-SUR-RHÓNE X (ARDÈCHE, FRANCE)
MILANO LUGLIO 2004
Elenco delle Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume I
I - CORNALI A E., 1 865 - Descrizione di una nuova specie dei genere Felis:
Felis jacobita (Corn.). 9 pp., I tav.
II - MAGNI-GR1FFI F, 1865 - Di una specie d 'Hippolais nuova per l'Italia.
6 pp., 1 tav.
Ili - GASTALDI B„ 1865 - Sulla riescavazione dei bacini lacustri per opera degli antichi ghiacciai. 30 pp.. 2 fìgg., 2 lavv.
IV - SEGUENZA G„ 1865 - Paleontologia malacologica dei terreni terzia-
rii del distretto di Messina. 88 pp.. 8 tavv.
V - GIBELLI G., 1865 - Sugli organi riproduttori del genere Verrucaria. 16
pp.. 1 tav.
VI - BEGG1ATO F. S., 1865 - Antracoterio di Zovencedo e di Monteviale
nel Vicentino, IO pp.. 1 tav.
VII - COCCHI t.,1865 - Di alcuni resti umani e degli oggetti di umana industria dei tempi preistorici raccolti in Toscana. 32 pp.. 4 tavv.
Vili -TARGION1TOZZETTI A. 1866 - Come sia fatto l'organo che fa lume nella lucciola volante dell' Italia centrale (Luciola italica ) e come le fibre muscolari in questo ed altri Insetti ed Artropodi. 28 pp.. 2 tavv.
IX - MAGGI L., 1865 - Intorno al genere Aeolosoma. 18 pp.. 2 tavv.
X - CORNALI A E., 1865 - Sopra i caratteri microscopici offerti dalle
Cantaridi e da altri Coleotteri facili a confondersi con esse. 40 pp.. 4 tavv.
Volume II
I - ISSEL A., 1866 - Dei Molluschi raccolti nella provincia di Pisa, 38 pp.
II - GENTILLI A., 1866 - Quelques considérations sur Forigine des bassins
lacustres, àpropos des sondages du Lac de Come. 12 pp.. 8 tavv.
Ili - MOLON F„ 1867 - Sulla flora terziaria delle Prealpi venete. 140 pp.
IV - D’ACHIARD! A., 1866 - Corallarj fossili del terreno nummulitico delle Alpi venete. 54 pp.. 5 tavv.
V - COCCHI L, 1866 - Sulla geologia dell'alta Valle di Magra. 18 pp.. 1 tav.
VI - SEGUENZA G., 1866 - Sulle importanti relazioni paleontologiche di
talune rocce cretacee della Calabria con alcuni terreni di Sicilia e dell’Africa settentrionale. 18 pp.. 1 tav.
VII - COCCHI I., 1866 - L'uomo fossile nell'Italia centrale. 82 pp.. 21 fìgg.. 4 tavv.
Vili - GAROVAGLIO S„ 1866 - Manzonia cantiana, novum Lichemmi Angiocarporum genus proposi timi atipie descriptum. 8 pp. 1 tav.
IX - SEGUENZA G., 1867 - Paleontologia malacologica dei terreni terzia¬
ri! del distretto di Messina (Pteropodi ed Eteropodi). 22 pp., 1 tav.
X - DURER B., 1867 - Osservazioni meteorologiche fatte alla Villa Carlotta
sul lago di Como, ecc. 48 pp. 11 tavv.
Volume III
I - EMERY C., 1873 - Studii anatomici sulla Vìpera Redii. 16 pp.. 1 tav.
II - GAROVAGLIO S., 1867 - Thelopsis. Betonici, Weitenwebera et
Limboria. qaatuor Lichemmi Angiocarporum genera recognita iconi- busque illustrata. 12 pp.. 2 tavv.
Ili - TARGIONI-TOZZETT1 A., 1867 - Studii sulle Cocciniglie. 88 pp.. 7 tavv.
IV - CLAPARÈDE E. R. e PANCERI P, 1867 - Nota sopra un Alciopide
parassito della Cydippe densa Forsk. 8 pp. 1 tavv.
V - GAROVAGLIO S., 1871 - De Pertusariis Eitropae medine commentatio.
40 pp.. 4 tavv.
Volume IV
I - D’ACHIARDI A., 1868 - Corallarj fossili del terreno nummulitico
dell’ Alpi venete. Parte 11 .32 pp. 8 tavv.
II - GAROVAGLIO S., 1868 - Ottona Lichenum genera ve! adhuc contro¬
versa. vel sedis prorsus incertae in svstemate. novis descriptionibus iconibusque accuratissimis illustrata. 18 pp., 2 tavv.
Ili - MARINONI C., 1868 - Le abitazioni lacustri e gli avanzi di umana industria in Lombardia. 66 pp.. 5 fìgg., 7 tavv.
IV - (Non pubblicato).
V - MARINONI C\, 1871 - Nuovi avanzi preistorici in Lombardia. 28 pp.,
3. fìgg-. 2 tavv.
NUOVA SF.RIF.
Volume V
I - MARTORELL1 G., 1895 - Monografia illustrata degli uccelli di rapina in Italia. 216 pp., 45 fìgg., 4 tavv.
Volume VI
I - DE ALESSANDRI G., 1897 - La pietra da cantoni di Rosignano e di
Vignale. Studi stratigrafici e paleontologici. 104 pp., 2 tavv., 1 carta.
II - MARTORELLI G. 1898 - Le forme e le simmetrie delle macchie nel
piumaggio. Memoria ornitologica. 112 pp., 63 fìgg., 1 tavv.
Ili - PAVESI P.. 1901- L'abbate Spallanzani a Pavia. 6# pp., 14 fìgg., 1 tav.
Volume VII
I - DE ALESSANDRI G., 1910 - Studi sui pesci triasici della Lombardia. 164 pp., 9 tavv.
Volume Vili
I - REPOSSI E., 1915 - La bassa Valle della Mera. Studi petrografici e geo¬
logici. Parte \. pp. 1-46, 5 fìgg.. 3 tavv.
II - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi petrogra¬
fici e geologici. Parte 11. pp. 47-186. 5 fìgg. 9 tavv.
Ili - AIRAGH1 C., 1917 - Sui molari d’elefante delle alluvioni lombarde, con osservazioni sulla filogenia e scomparsa di alcuni Proboscidati. pp. 187-242, 4 fìgg., 3 tavv.
Volume IX
I - BEZZI M. 1918 - Studi sulla ditterofauna nivale delle Alpi italiane, pp.
1-164. 7 fìgg. 2 tavv.
II - SERA G. L., 1920 - Sui rapporti della conformazione della base del cra¬
nio colle forme craniensi e colle strutture della faccia nelle razze umane. (Saggio di una nuova dottrina craniologica con particolare riguardo dei principali cranii fossili)./;/;. 165-262. 7 fìgg.. 2 tavv.
Ili - DE BEAUX O. e FESTA E., 1927 - La ricomparsa del Cinghiale nell'Italia settentrionale-occidentale, pp. 263-320, 13 fìgg.. 7 tavv.
VolumeX
I - DESIO A., 1929 - Studi geologici sulla regione dell’Albenza (Prealpi
Bergamasche)./;/;. 1-156. 27 fìgg.. I tav, 1 carta.
II - SCORTECCI G., 1937 - Gli organi di senso della pelle degli Agamidi.
pp. 157-208, 39 fìgg. 2 tavv.
Ili - SCORTECCI G.. 1941-1 recettori degli Agamidi. pp. 209-326, 80 fìgg.
Volume XI
1 - GU1GLIA D„ 1944 - Gli Sfecidi italiani del Museo di Milano (Hvmen.). pp. 1-44. 4 fìgg.. 5 tavv.
11-111 - G1ACÓM1N1 V. e PIGNATTI S., 1955 - Flora e Vegetazione dell'Alta Valle del Braulio. Con speciale riferimento ai pascoli di alti¬ tudine. /;/;. 45-238, 31 fìgg., 1 carta.
Volume XII
1 - VIALL1 V., 1956 - Sul rinoceronte e l'elefante dei livelli superiori della serie lacustre di LefTe (Bergamo)./;/;. 1-70. 4 fìgg. 6 tavv.
I -VENZO S., 1957 - Rilevamento geologico dell'anfiteatro morenico del Garda. Parte 1: Tratto occidentale Gardone-Descnzano. pp. 71-140, 14 fìgg.. 6 tavv, 1 carta.
Ili - V1ALLI V., 1959 - Ammoniti sinemuriane del Monte Albenza (Bergamo)./;/;. 141-188, 2 fìgg.. 5 tavv.
Volume XIII
I - VENZO S., 1961- Rilevamento geologico dell'anfiteatro morenico del
Garda. Parte II. Tratto orientale Garda-Adige e anfiteatro atesino di Rivoli veronese./;/;. 1-64, 25 fìgg., 9 tavv., 1 carta.
II - PINNA G., 1963 - Ammoniti del Lias superiore (Toarciano) delFAlpe
Turati (Erba, Como). Generi Mercaticeras, Pseudomercaticeras e Brodieia. pp. 65-98, 2 fìgg.. 4 tavv.
Ili - ZANZUCCHI G., 1963 - Le Ammoniti del Lias supcriore (Toarciano) di Entratico in Val Cavallina (Bergamasco orientale). /;/;. 99-146. 2 fìgg. 8 tavv.
Volume XIV
I - VENZO S., 1965 - Rilevamento geologico dell’anfiteatro morenico fron¬
tale del Garda dal Chiese all'Adige, pp. 1-82, 1 ffìgg-, 4 tavv, I carta.
II - PINNA G„ 1966 - Ammoniti del Lias superiore (Toarciano) dell’Alpe
Turati (Erba, Como). Famiglia Dactvlioceratidae. pp. 83-136, 4 tavv.
Ili - DIEN1 I.. MASSARI F. ^MONTANARI L„ 1966 - Il Paleogene dei dintorni di Orosei (Sardegna)./;/;. 13-184, 5 fìgg., 8 tavv.
Volume XV
I - CARETTO P. G., 1966 - Nuova classificazione di alcuni Briozoi plioce¬
nici, precedentemente determinati quali Idrozoi del genere Hydractinia Vati Beneden. pp. 1-88, 27 fìgg. 9 tavv.
II - D1EN1 1. e MASSARI F„ 1966 - Il Neogene e il Quaternario dei dintor¬
ni di Orosei (Sardegna), pp. 89-142, 8 fìgg.. 7 tavv.
Ili - BARBIERI F„ IACCARINO S„ BARBIERI F. & PETRUCC1 F., 1967 - Il Pliocene del Subappennino Piacentino-Parmense-Reggiano. pp. 143-188, 20 fìgg., 3 tavv.
Volume XVI
I - CARETTO P. G., 1967 - Studio morfologico con l'ausilio del metodo sta¬
tistico e nuova classificazione dei Gasteropodi pliocenici attribuibili al Murex brandaris Linneo . pp. 1-60, 1 fìg., 7 tabb., 10 tavv
II - SACCFH VIALLI G. e CANTALUPPI G„ 1967 - I nuovi fossili di
Gozzano (Prealpi piemontesi)./;/;. 61-128, 30 Jìgg., 8 tavv.
Ili - P1GORINI B., 1967 - Aspetti sedimentologici del Mare Adriatico, pp. 129-200, 13 fìgg., 4 tabb. 7 tavv.
Volume XVII
I - PINNA G.. 1968 - Ammoniti del Lias superiore (Toarciano) dell'Alpe Turati (Erba, Como). Famiglie Lytoceratidae, Nannolvtoceralidae.
Anna Alessandrello, Giacomo Bracchi & Bernard Riou
Museo Civico di Storia Naturale di Milano Musée de Paléontologie de la Voulte
Polychaete, sipunculan and enteropneust worms from thè Lower Callovian (Middle Jurassic) of La Voulte-sur-Rhòne (Ardèche, France)
Volume XXXII - Fascicolo I
Luglio 2004
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
INDEX
Introduction
Geological Setting and Preservation
Systematic Paleontology
Unrelated specimens
Discussion and Conclusions
Acknowledgements
References
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© 2004 Società Italiana di Scienze Naturali e Museo Civico di Storia Naturale Corso Venezia, 55 - 20121 Milano
In copertina: Paleoaphrodite gallica. Disegno di Fabio Fogliazza
Registrato al tribunale di Milano al n. 6694
Direttore responsabile: Anna Alessandrello Responsabile di redazione: Stefania Nosotti Grafica editoriale: P.R.G.
Stampa: Litografia Solari, Peschiera Borromeo - Giugno 2004
ISSN 0376-2726
Anna Alessandrello, Giacomo Bracchi & Bernard Riou
Polychaete, sipunculan and enteropneust worms from thè Lower Callovian (Middle Jurassic) of La Voulte-sur-Rhòne (Ardèche, France)
Abstract - The worm fauna from thè Lower Callovian (Middle Jurassic) of La Voulte-sur-Rhòne (Ardèche, France) is described through thè recog- nition of two new genera and five new species of polychaete annelids: thè arenicoloidean lubarenicola fìscheri n. gen. n. sp. and thè aphroditoideans Paleoaphrodite gallica n. sp., Paleoaphrodite briggsiana n. sp., Paleoaphrodite adeliae n. sp. and Protopholoe rhodanitidis n. gen. n. sp. Moreover, Rondeletia scutata n. gen n. sp. is thè third sipunculan species recorded within Mesozoic, Megaderaion callovianum n. sp. is thè third fossil enterop¬ neust species known. Since thè soft tissues of these worms show different degrees of pyritization, some considerations about thè La Voulte-sur-Rhòne paleoenviroment are proposed.
Key words - Mesozoic, France, polychaetes, sipunculans, enteropneusts, pyritization.
Riassunto - Gli organismi vermiformi del Calloviano inferiore (Giurassico medio) di La Voulte-sur-Rhòne (Ardenne, Francia).
Vengono descritti gli organismi vermiformi del giacimento calloviano (Giurassico medio) di La Voulte-sur-Rhòne (Ardenne, Francia). Due nuovi generi e cinque nuove specie di annelidi policheti sono riconosciuti: lubarenicola fìscheri n. gen. n. sp. (ordine Capitellida), Paleoaphrodite gallica n. sp., Paleoaphrodite briggsiana n. sp., Paleoaphrodite adeliae n. sp. e Protopholoe rhodanitidis n. gen. n. sp. (superfamiglia Aphroditoidea). Sono inol¬ tre descritti la terza specie di sipunculide conosciuta per il Mesozoico, Rondeletia scutata n. gen n. sp., e il terzo enteropneusto fossile conosciuto, Megaderaion callovianum n. gen. n. sp. Essendo i tessuti molli degli esemplari allo studio conservati attraverso vari gradi di piritizzazione, è stato infi¬ ne possibile proporre alcune considerazioni sull’ambiente di deposizione.
Parole chiave - Mesozoico, Francia, policheti, sipunculidi, enteropneusti, piritizzazione.
INTRODUCTION
Fossil worms are rare in Mesozoic Lagerstàtten. In addi- tion to thè eunicemorph polychaetes from thè Kimmeridgian (Upper Jurassic) of Solnhofen, Germany (Ehlers, 1 868 and 1 869), and from thè Buntsandstein (Lower Triassic) of Grès a Voltzia, France (Gali & Grauvogel, 1 966), fossil polychaetes, nematodes and enteropneusts are known from thè Sinemurian (Lower Jurassic) of Osteno, northem Italy (Arduini et al., 1981, 1982 and 1983). Triassic fossil polychaetes have been reported also from thè Rhaetian (Lower Triassic) of thè Italian Prealps (Alessandrello & Teruzzi, 1986a; Stefani et al., 1992) and from thè Olenekian (Lower Triassic) of thè Ambilobè region, Madagascar (Alessandrello, 1990), represented by aphroditid species. Triassic worms also include an unrelated species from thè Camian of Slovenia (Kolar-Jurkovsek & Jurkovsek, 1997). However, thè most diverse and abundant Mesozic worm fauna has been described recently from thè Middle Cenomanian (Upper Cretaceous) of Lebanon, includ- . ing 6 families, 7 genera and 1 6 species of polychaete annelids (Alessandrello & Teruzzi, 1986b; Bracchi & Alessandrello, 2004, in press) and a new genus of a tetrameric nematode (Alessandrello & Mattavelli, 2004, in press).
The Callovian outcrop of La Voulte-sur-Rhòne is wide- ly known for thè exceptional preservation of some inverte¬ brate taxa such as bivalves, gastropods, cephalopods, pyc-
nogonids, ophiuroids and asteroids (Fischer, 2003). Thylacocephalan, macruran, cumacean and mysidacean crustaceans have been recorded also (Van Straelen, 1922 and 1923; Secretan & Riou, 1983 and 1986; Carriol & Riou, 1991). Among thè vertebrates, a few elasmobranchs, actinopterygians and crossopterygians (Fischer, 2003) have been described together with a crocodilian skull (Kuhn- Schnyder, 1960). Moreover, in thè last thirty years, some authors (Dietl & Mundlos, 1972; Briggs & Kear, 1993; Fischer, 2003) have mentioned thè existence, in thè La Voulte-sur-Rhòne outcrop, of worms in generic terms referred to aphroditid and arenicolid polychaetes. In this paper we describe 23 worm specimens from thè fìiveau e ’ of La Voulte-sur-Rhòne (Fischer, 2003), held by thè collec- tions of thè Laboratoire de Paléontologie (L.P.M.) of thè Muséum National d’Histoire Naturelle in Paris.
The rock slabs preserving thè La Voulte-sur-Rhòne worms have been analyzed through a x-radiographic set (Fig. 1). This method could have been useful to guide preparation of thè fossils but did not yield information beyond that directly observable through thè microscopi- cal investigation.
The polychaete systematics adopted in this work fol- lows Fauchald (1977).
4
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
Fig. 1 - X-radiographs of L.PM.-B.48343 and L.P.M.-B.48353 (A and B, Iubarenicola fischeri n. gen. n. sp.), L.P.M.-B.48336 (C, Paleoaphrodite brig- gsiana n. sp.), I PM.-B.48340 (D, Pmtopholoe rhodanitis n. gen. n. sp.), L..P.M. B. 48345 (E, Rondeletia scutata n. gen. n. sp.) and L.P.M.-B.48347 (F, undetermined worm).
POLYCHAETE, SIPUNCULAN AND ENTEROPNEUST WORMS FROM THE LOWER CALLOVIAN
5
GEOLOGICAL SETTING AND PRESERVATION
The La Voulte-sur-Rhòne fossiliferous levels take their name from a small town on thè west bank of thè Rhone River, in thè French southeastern department of Ardèche, and outcrop at La Boissine, a locality along thè northern banks of thè Gramade stream, slightly west of thè La Voulte-sur-Rhòne iron mines (Etter, 2002a; Fischer, 2003). The La Voulte-sur-Rhòne stratigraphy has been investigated in detail by Elmi (1967), who dated this outcrop to thè Lower Callovian on thè basis of thè ammonite fauna ( gracilis biozone, koenigi sub-biozone). Argillaceous and finely laminated marlstones showing a black, brown, reddish or variegated colour and varying in thickness from 3 m to 7 m, alternate discontinuously with rare calcareous strata. Apart from widespread calcareous, siliceous and pyritic nodules, thè marly levels often con- tain limonite and siderite traces and concretions.
The La Voulte-sur-Rhòne fossil worms show different kinds of preservation. That described in some cephalopods (Fischer & Riou, 1982a and 1982b; Etter,
2002a) is surely thè most spectacular: soft parts are pre- served as pyritized three-dimensional relief. This is also thè case for thè aphroditoideans and thè enteropneust described here. However, as already pointed out for thè other fossil remains from La Voulte-sur-Rhòne (Etter, 2002a; Fischer, 2003), some specimens are preserved as organic traces in a calcareous or marly-argillaceous matrix, fossilized through substitution by limonite, pyrite or calcite mixed with some argillaceous elements. According to Wilby et al. (1995 and 1996) and Wilby (2001), thè La Voulte-sur-Rhòne soft tissues preserved in thè marly-argillaceous matrix and those preserved in early diagenetic concretions underwent a different kind of substitution. In thè first case they are flattened on thè rock slab and largely made up of apatite, while in thè second they are three-dimensional reliefs in which calcite, quartz, gypsum and pyrite are associated in thè diagenet¬ ic sequence that preserved thè organism.
SYSTEMATIC PALEONTOLOGY
Phylum Annelida Lamarck, 1 809 Class Polychaeta Grube, 1850 Order Capitellida Fauchald, 1977 Family uncertain Genus Iubarenicola n. gen.
Diagnosti: as for thè type species.
Etymology : from thè Latin iuba (=crest) and from Arenicola , a living arenicoloidean genus.
Type species : Iubarenicola fìscheri n. sp.
,
Iubarenicola fìscheri n. sp.
Fig. 2, PI. 1A-B
Diagnosti : elongate body comprised of several short metameres of variable length. Two mediai, thin, parallel ridges on thè dorsal surface delimit a shallow furrow from thè first metamere behind thè prostomium to thè pygidi- um. Prostomium is small and subconical.
Etymology : dedicated to Jean-Claude Fischer, in recognition of his studies about thè La Voulte-sur-Rhòne fossils.
Geological age: Lower Callovian (Middle Jurassic).
Type locality: La Boissine, La Voulte-sur-Rhòne.
Material: 2 specimens. L.P.M.-B.48343 (holotype, Fig. 2A) and L.P.M.-B.48353 (paratype, Fig. 2B).
Description: both thè holotype and paratype are pre¬ served as a three-dimensional pyritized relief. In thè holo¬ type, thè body width gradually decreases fforn thè pros- '. tomium to thè pygidium. The paratype body is narrow both at thè proximal and distai extremities and it shows a slight widening in thè posterior half. Both thè holotype and paratype are about 65 mm long and they appear coiled in thè posterior part. The holotype is preserved as a dorsoventral compression and thè dorsal surface is exposed. The ventral side of thè body is flattened and thè
dorsal is convex: it is difficult to distinguish if this is a preservation artifact or a reai feature. The paratype under¬ went twisting, so that thè dorsal and ventral surfaces are evident for different body portions. The body shows a clear metamerical organization, but only in thè holotype can all thè metameres be counted: there are 65 metameres each about 1 mm long.
The mediai ridges developed along thè dorsal surface are particularly evident in thè anterior part of thè body, but they are well preserved in other sections also. It is impossible to establish if thè axial furrow reflects a spe¬ cial internai morphology.
The cephalic extremity of thè paratype is connected to a sub-circular and slightly flattened structure sensibly wider than thè body: we interpret this structure as thè glo¬ bose everted pharynx typical of some Capitellida (Fauvel, 1927; Fauchald, 1977).
Chaetae (setae) are not preserved.
Remarks: thè generai body morphology together with thè little sub-conical prostomium and thè presence of a globose everted pharynx suggest to assign thè examined specimens to thè order Capitellida or to thè order Orbinida Fauchald, 1977. The Orbinida can own a pros¬ tomium very similar to that of Capitellida, but their ever¬ sale pharynx appears as an axial sac or a ventrolateral pad. However, thè examined samples lack other charac- ters that would allow us to determine a more precise posi- tion within Capitellida.
Capitellida are known from thè Triassic to Recent (Howell, 1962). Archarenicola rhaetica Horwood, 1912 from thè Rhaetian (Lower Triassic) of England is charac- terized by a frilled prostomium. The genus Arenicola Lamarck, 1801 has been recorded from thè Cretaceous on thè basis of trace fossils (Martinell et al ., 2001).
6
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
Fig. 2 - lubarenicola fìscheri n. gen. n. sp., holotype (L.P.M.-B.48343, A, x 1.5) and paratype (L.P.M.-B. 48353, B, x 1.5).
Order Phyllodocida Dales, 1962 Superfamily Aphroditoidea Malmgren, 1 867 Family Aphroditidae Malmgren, 1 867 Genus Paleoaphrodite Alessandrello & Teruzzi, 1986 Paleoaphrodite gallica n. sp.
Fig. 3, PI. 1C
Diagnosti: small-sized and slightly elongate fusiform aphroditid. Biramous parapodia feature two series of overlapped, crossed and posteriorly oriented setal tufìts of variable length and thickness. Each tuft is made up of two setal types.
Etymology : from thè Latin gallicus (=belonging to thè Roman province of Gaul), referring to thè geographical origin.
Geological age : Lower Callovian (Middle Jurassic).
Type locality : La Boissine, La Voulte-sur-Rhòne.
Material : 5 specimens. L.P.M.-B.48342 Aa (part and counterpart, holotype, Figs. 3A-3B), L.P.M.-B.48341 Aa- g (paratype, Fig. 3C), L.P.M.-B.48337 Ae, L.P.M.- B. 48338 Ae (part and counterpart) and L.P.M.-B. 48339 Aa-g (part and counterpart).
Description : all thè examined specimens are complete and measure from 18 mm (L.P.M. B48337 Ae) to 34 mm (1 P.M. 48342 Aa) in length. The maximum width of thè body is 5 mm, excluding chaetae, in all thè specimens.
Chaetae are clumped in two series of tufts on each side of thè body, clearly indicating biramous parapodia. The tufts of adjacent metameres overlap, as it is clearly evi-
dent in L.P.M. 48342 Aa and L.P.M.-B.48341 Aa-g. The maximum length of thè chaetae grades from 5 mm to 2 mm. On each side of thè body, chaetae form on average 20 pairs of tufts (24 in L.P.M.-B.48341 Aa-g, 18 in L.P.M.-B. 48342 Aa) disposed in two overlapping series: thè chaetae belonging to thè upper series are pressed down on those of thè lower series, so as it is impossible to distinguish between thè notopodial and neuropodial ones. Each tuft comprises an undetermined number of very thin and subcylindrical chaetae, measuring less than 0.1 mm in thickness, associated with two or three stronger chaetae, maybe aciculae, measuring about 0.1 mm in thickness. The junction between setal tufts and body is only clearly evident in two tufts near thè mid-length of L.P.M.-B.48341 Aa-g, because in thè other specimens only thè mediai and distai parts of thè chaetae are pre- served. However, in these other specimens some attach- ment points are recognizable as little depressions (in parts) or protuberances (in counterparts) at thè proximal extremity of thè tufts.
The metamerical subdivision of thè body is clearly recognizable in L. P.M. -B. 48342 Aa, where 10 segments are preserved near thè caudal extremity. Each metamere is 1 mm long and appears to carry two pairs of setal tufts. In thè mid-part of thè body of L.P.M.- B. 48342 Aa, five metameres are distinguished by pro- nounced constrictions.
Remarks : thè generai body morphology allows thè exam¬ ined material to be assigned to thè superfamily Aphroditoidea
POLYCHAETE, SIPUNCULAN AND ENTEROPNEUST WORMS FROM THE LOWER CALLOVIAN
7
Fig. 3 - Paleoaphrodite gallica n. sp., holotype (L.P.M.-B. 48342 Aa: A, part, x 4.4; B, counterpart, x 3) and paratype (L.P.M.-B.48341 Aa-g, C; x 2.8).
and, together with thè arrangement of chaetae in two tufts that define thè margins of each metaniere, suggest a relationship with thè family Aphroditidae and thè genus Paleoaphrodite. To date two Mesozoic aphroditids have been assigned to Paleoaphrodite. However, in contrast to P gallica n. sp., P raetica Alessandrello & Teruzzi, 1 986 from thè Rhaetian of Lombardy (northem Italy) shows an ovai body. Moreover, in P. gallica n. sp. thè two tufts of each parapodium are com- posed of both strong and thin chaetae, whereas in P raetica thè strongest chaetae occur both in thè upper and lower tufts, while thè thinnest are limited to thè lower ones.
Among thè species of Paleoaphrodite already described, P anaboranoensis Alessandrello, 1990 from thè Lower Triassic of Madagascar has a fusiform body, but it is shorter and wider than that of P. gallica n. sp. Moreover, thè two pairs of setal tufts that occur on each metamere of P. anaboranoensis are made up of strong chaetae only. P. brig- gsiana n. sp. and P. adeliae n. sp. also show a fusiform body, but their larger dimensions, together with thè pres- ence of a jaw apparatus and a different chaetae architecture, distinguish them from P. gallica n. sp.
Paleoaphrodite briggsiana n. sp.
Fig. 4, PI. 1D
Diagnosis : elongate fusiform body. Because of thè metamere spacing, setal tufts of each metamere do not over- lap nor traverse adjacent ones. Each tuft comprises subcylin- drical and flattened chaetae. Pharynx armoured with two pairs of jaws. Very large and semicircular prostomium.
8
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
Etymology : dedicateci to Derek E.G. Briggs, in recog- nition of his taphonomic studies of thè La Voulte-sur- Rhòne fossils.
Geological age : Lower Callovian (Middle Jurassic).
Type locality : La Boissine, La Voulte-sur-Rhòne.
Material : 1 specimen. L.P.M.-B.48336 (holotype).
Description : thè specimen is preserved as a three- dimensional pyritized relief measuring 67 mm in length: only thè dorsal side is evident. The maximum width excluding chaetae occurs near thè mid-length: 1 8 mm.
The prostomium is very large and semicircular, meas¬ uring at least 10 mm in length and 25 mm in width.
Even if thè poor presevation of L.P.M.-B.48336 does not allow a detailed description, at least 14-16 protuber- ances, that can be interpreted as thè attachments of thè upper series of setal tufts, are clearly evident on each side of thè body. At thè mid-length and in thè posterior pari of thè body, thè shape and position of thè protuberances sug- gest a reai metamerism, evidenced by thè presence of 3 segments separated by thin furrows.
The chaetae preserved on each side of thè body are organized in tufts, separated from those of thè adjacent metameres by 2-3 mm gaps. Neither thè originai number nor thè length of thè chaetae can be determined, and thè biramous structure of thè parapopodia is obscured. However, each tuft appears to consist of two types of strong chaetae: some chaetae, maybe aciculae, show a cylindrical shape, a yellowish or whitish semitransparent colour and a width of 0.1 mm; other chaetae show thè same width but are more flattened and darker in colour. It is impossible to establish whether both kinds of chaetae were present in thè same tuft of each parapodium or not.
Near thè anterior extremity of thè body thè elements that make up thè jaw apparatus are raised slightly above thè pyritized relief of thè body as little black and shiny structures, but it is very difficult to distinguish their organization and shape; some elements may have been displaced from their former position. Elements here pre¬ served are two scythe-shaped jaws, showing backward oriented tips and converging anteriorly toward a strong semicircular piate, and two smaller and thinner hook- shaped jaws. These last appear to be linked posteriorly, but probably they were formerly separated and arranged on opposite sides of thè jaw. The semicircular piate is not in its originai position: it does not lie in thè same piane as thè jaws, but is oriented perpendicular to thè bedding piane. There is no evidence that thè pharynx is everted, but fossilization of thè jaws may be more likely when thè pharynx is everted: this is because thè La Voulte-sur- Rhòne outcrop is characterized by rapid pyritization of thè soft tissues after death (Wilby et al., 1996), so that thè presence of formerly mineralized tissues, such as poly- chaete jaws, may be concealed. The hypothesis that thè P.M.-B.48336 pharynx is everted is supported by thè 'b ervation that it is thè only polychaete specimen from toissine in which thè jaw apparatus is recognizable near its originai position.
Ree rks: thè generai body morphology allows thè specimen to be assigned to thè family Aphroditidae and to thè genus Paleoaphrodite. However, thè setal architec- ture, thè metaniere development and thè presence of an
armoured pharynx, allow P. briggsiana n. sp. to be distin- guished from thè previously described species in thè genus. In addition to P. briggsiana n. sp., thè other Paleoaphrodite species owning a fusiform body are P. gallica n. sp. and P adeliae n. sp.. P adeliae n. sp. is very similar in body shape, but it is very different in thè devel¬ opment of thè prostomium as well as in thè jaw apparatus structure. The fusiform body of P gallica n. sp. is sensi- bly more elongate and narrow if compared with that of P briggsiana n. sp. and P adeliae n. sp., in which is more evident a sensible tapering toward both thè cephalic and caudal extremities. With regard to thè setal organization P adeliae n. sp. is very similar to P briggsiana n. sp., because of thè presence of both subcylindrical and flat¬ tened chaetae: however, in P briggsiana n. sp. thè setal tufts do not form a continuous coat around thè body, but they appear separated by 2-3 mm gaps produced by thè major spacing among metameres. Differently from both P briggsiana n. sp. and P adeliae. n. sp., P gallica n. sp. owns subcylindrical chaetae only.
Among thè fossil aphroditids described to date, thè presence of a pharyngeal armature including two pair of jaws has been observed only in thè aphroditids of thè sub- family Hystriciolinae Thompson, 1979, Dryptoscolex matthiesae Thompson, 1979 and Fastuoscolex gemmatus Thompson, 1979, both from thè Carboniferous of Mazon Creek (Thompson, 1979): this is probably due both to thè absence of jaws in some aphroditid species and to arti- faets of preservation.
Fig. 4 - Paleoaphrodite briggsiana n. sp., holotype (L.P.M.-B. 48336, x 0.8).
POLYCHAETE, SIPUNCULAN AND ENTEROPNEUST WORMS FROM THE LOWER CALLOVIAN
9
Paleoaphrodite adeliae n. sp.
Fig. 5, PI. 1E
Diagnosti: large fusiform body. Two setal types forming a continuous coat around thè body and increasing in length from prostomium to pygidium. Jaw apparatus present.
Etymology : dedicated to Adelia Bracchi, for her strong support in recent years.
Geological age : Lower Callovian (Middle Jurassic).
Type locality : La Boissine, La Voulte-sur-Rhòne.
Material : 1 specimen. L.P.M.-B.48335 (holotype).
Description : excluding thè setal coat, thè body of thè holotype measures 5 1 mm in length and 1 7 mm in width. It is preserved as a three-dimensional pyritized relief revealing thè dorsal side of thè worm. The subtriangular prostomium measures 4 mm in length and 8 mm in width.
Along each side of thè body a single series of aligned protuberances is evidenti they probably represent thè attachment points of thè upper series of setal tufts. At least 20 protuberances are evident along thè left side of thè body, at least 15 along thè right side.
Two kinds of chaetae are present: some chaetae, maybe aciculae, are about 0.1 mm wide and they show a subcylindrical shape and a semitransparent yellowish or whitish colour, while others are slightly thinner, flattened and dark coloured. The chaetae have broken along their length when thè slab was split, so it is impossible to deter¬ mine their exact length or whether they are notopodial or neuropodial, although it is evident that their length increases from thè cephalic to thè caudal extremity. The poor quality of preservation obscures thè biramous nature of thè parapodia. The chaetae of thè anterior part of thè body are tumed forward, whereas those of thè posterior part are tumed backward.
The protuberances clearly indicate thè metamerical subdivision of thè body. The metameres decrease in length from thè mid part of thè body (0.3 mm) toward both thè anterior and posterior extremity. The first metameres are concave anteriorly, but they become con- vex toward thè caudal extremity of thè body. The furrows that separate thè metameres are evident only in thè last quarter of thè body.
The body is covered by several thin, shallow and lon- gitudinally developed furrows: they probably represent thè imprint of thè felt of capillary fibers that typically covers thè dorsal surface of some aphroditids. These fibers are observed here for thè first time in Paleoaphrodite : they are, however, considered poorly diagnostic because their presence or absence could be thè result of differences in preservation.
Near thè body center, a small, black, shiny, markedly mineralized and hook-shaped structure hearing a strong tooth near thè anterior extremity of its convex side is clearly evident: it is one of thè elements comprising thè pharyngeal armature and it is slightly displaced fforn its originai position.
Remarks: among described species of Paleoaphrodite , that most similar to P adeliae n. sp. is P. briggsiana n. sp., particularly in generai body morphology, shape and setal organization. Moreover, subcylindrical and semitranspar¬ ent chaetae identical to those of P adeliae n. sp. are also
present in P briggsiana n. sp.. However, thè latter has also flattened chaetae. Other dilferences between these two species are thè architecture of thè jaws, thè shape of thè prostomium and thè organization of thè setal tufts: in P adeliae n. sp. thè tufts of adjacent metameres overlap, forming a continuous coat around thè whole body, and thè prostomium is subtriangular. In P briggsiana n. sp. thè tufts of adjacent metameres are clearly separated and thè prostomium is larger and semicircular.
Fig. 5 - Paleoaphrodite adeliae n. sp., holotype (L.P.M.-B. 48335, x 1.7).
Family uncertain Genus Protopholoe n. gen.
Diagnosti: as for thè type species.
Etymology: from thè Greek n-pcÓTos (=primitive) and from Pholoe, living genus.
Type species: Protopholoe rhodanitis n. sp.
Protopholoe rhodanitis n. sp.
Fig. 6, PI. 1F
Diagnosti: dorsally convex, short and subquadrangu- lar body with several metameres and a deep, straight, nar- row and slightly concave furrow developed along thè median body line. Slightly elongate and ovai prostomium. Short and strong aciculae.
Etymology: from thè Latin rhodanitis (=belonging to Rhodanus ), referring to thè Rhone Valley.
Geological age: Lower Callovian (Middle Jurassic).
Type locality: La Boissine, La Voulte-sur-Rhòne.
Material: 1 specimen. L.P.M.-B. 48340 (holotype).
10
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
Description : holotype preserved as a three-dimension- al pyritized relief measuring 13 mm in length: only thè dorsal side is evident. The body width varies from 2 mm at thè cephalic and caudal extremities to 4 mm in thè cen¬ trai part. Twenty metameres separated by shallow furrows are clearly evident. The first seven metameres are con¬ cave anteriorly, a condition becoming less pronounced from prostomium to pygidium. The eight longest metameres are those of thè centrai body section: in some cases they are twice thè length of those at thè cephalic and caudal extremities.
Some posteriorly directed aciculae are preserved as mineralized structures along thè left side of thè body only, but their parapodial arrangement is not evident.
Remarks : thè generai body morphology supports an assignment of thè specimen examined to thè superfamily Aphroditoidea. In particular, thè specimen shows strong similarities with thè modern genus Pholoe Johnston, 1839, which is actually included in Sigalionidae Malmgren, 1867 (Glasby et al., 2000), soft bottoni scale- worms that ofìten show a subquadrangular and flattened body (Fauchald, 1977; Fauchald & Rouse, 1997). Moreover, as in thè fossil, thè body of thè living genus Pholoe is dorsally convex and shows a deep centrai fur- row separating it into two symmetrical parts from thè cephalic to thè caudal extremity (Fauvel, 1923). However, because of thè absence of soft parts in thè fossil speci¬ men, further comparisons are not possible and thè sys- tematic position of P rhodanitis n. sp. within thè aphrodi- toideans is uncertain.
Fig. 6 - Protopholoe rhodanitis n. gen. n. sp., holotype (LPM- B.48340, x 4.5).
Phylum Sipuncula Stephen, 1935 Class Sipunculidea E.Cutler & Gibbs, 1994 Order and Family uncertain Genus Rondeletia n. gen.
Diagnosti: as thè type species.
Etymology: dedicated to Guidarne Rondelet, who first described and illustrated sipunculan worms during thè sixteenth century.
Type species : Rondeletia scutata n. sp.
Rondeletia scutata n. sp.
Fig. 7, PI. 1G-H-I
Diagnosti : slightly elongate body divided in two dis- tinct regions. A narrow region characterized by few longi- tudinal ridges is followed by a broader region characterized by several transverse ridges and equipped with a conical shield at its distai extremity. Body wall including some sep¬ arated and not anostomosing longitudinal muscle bands.
Etymology : from thè Latin scutatus (=equipped with a shield), referring to thè presence of a conical shield cov- ering thè caudal extremity.
Geological age: Lower Callovian (Middle Jurassic).
Type locality: La Boissine, La Voulte-sur-Rhóne.
Material : 6 specimens. L.P.M.-B.48355 and L.P.M.- B.48356 (holotype, part and counterpart, Figs. 7A-7B), L.P.M.-B.48344 (paratype, Fig. 7C), L.P.M.-B.48345, L.P.M.- B.48354 Aa, L.P.M.-B.48357 Aa and L.P.M.-B.48358 Aa.
Description : thè holotype is preserved as a complete external mold, while thè paratype is preserved as a three- dimensional pyritized relief, as are L.P.M.-B. 48345, L.P.M.-B. 48357 Aa and L.P.M.-B.48358 Aa, while L.P.M.-B.48354 Aa is preserved as imprint.
The body is completely preserved in L.P.M.-B.48354 Aa and in thè holotype only, where it appears rather coiled up. In both these specimens two distinct regions are evi¬ dent: in thè holotype thè narrower region is 30 mm long and characterized by thè presence of four longitudinal and par- allel ridges. This region decreases slightly in width from its posterior extremity (10 mm in holotype, 8 mm in L.P.M.- B.48354 Aa) toward thè anterior extremity. We interpret this region as an everted introvert, thè ridges as thè longitu¬ dinal muscle bands of thè body wall. In L.RM.-B.48354 Aa thè diagenetical compression of thè early mineralized body resulted in a slight widening of thè introvert, so that it appears, in contrast to thè other specimens, wider than thè rest of thè body: it is about 30 mm long.
The introvert is normally followed by a slightly wider region, here interpreted as thè trunk, that reaches a maxi¬ mum width near its mid-length and shows a slight tapering toward both its extremities. In thè holotype thè trunk meas- ures about 80 mm in length and 1 8 mm in maximum width, while in thè paratype, which only preserves thè trunk, thè length is 60 mm and thè maximum width 8 mm. For L.RM.-B.48354 Aa thè trunk length is 45 mm and thè max¬ imum width 10 mm. In L.P.M.-B.48357 Aa and L.P.M.- B.48358 only a fragment of thè anterior part of thè trunk is preserved, and in L.P.M.-B. 48345 thè trunk is completely preserved but it cannot be measured precisely, even if sim- ilar in dimension to that of thè holotype, because it is com¬ pletely coiled up. Particularly in thè paratype, thè trunk appears divided into several short but not equally developed segments, evidenced by transverse parallel ridges separated by shallow furrows. These ridges and furrows are probably
POLYCHAETE, SIPUNCULAN AND ENTEROPNEUST WORMS FROM THE LOWER CALLOVIAN
11
Fig. 7 - Rondeletia scutata n. gen. n. sp., holotype (L.P.M.-B.48355, A, part, x 1.6; L.P.M.-B. 48356, B, counterpart, x 1.6) and paratype (L.P.M.-B.48344, C, x 1.8).
evidence of pseudometamerism that affected thè cuticle only, supporting a sipunculan affinity. In thè paratype, L.P.M.-B.48354 Aa and in L.P.M.-B. 48358 Aa thè trunk also preserves two or three longitudinal ridges that we interpret in thè same way as those observed on thè introvert. In our opinion thè longitudinal ridges are more numerous and always evident on thè introvert because of its fragility, which was surely more marked than that of thè cuticle cov- ering thè trunk. In thè examined specimens, apart from thè cases described above, thè longitudinal ridges are limited to thè narrower portion of thè body and abruptly terminated where thè trunk widens: this allows thè introvert-trunk boundary to be identified.
The cuticle covering thè distai extremity of thè trunk shows no evidence of segmentation, but seems to form a subconical shield carrying a thin ridge along its median line that is clearly evident in thè holotype, paratype and L.P.M.-B.48354 Aa. The maximum dimensions of this shield are those in thè holotype, where it measures 7 mm in length and 8 mm in maximum width, while in thè paratype it is 4 mm long and 3 mm wide.
There are no traces of appendages.
Remarks: thè two distinct regions (introvert and trunk) that comprise thè body of thè examined specimens, together with thè presence of a caudal shield and of thè longitudinal muscle bands, justify their assignment to thè phylum Sipunculida. This phylum includes thè classes Phascolosomatidea E.Cutler & Gibbs, 1995 and Sipunculidea, identified on thè basis of several soft part features, usually not recognizable in fossils (Cutler, 1994), such as thè arrangement of thè tentacles surround- ing thè mouth at thè tip of thè introvert (Maxmen et al .,
12
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
2002). However, thè assignment of R. scutata n. gen. n. sp. to Sipunculidea is reasonably secure because within Phascolosomatidea only thè Aspidosiphonidae Baird, 1868 have a caudal shield but, in contrast to Sipunculidea, generally have an introvert protruding at an angle of 45- 90° to thè main axis of thè trunk and thè longitudinal muscle layers of thè body wall are separated into anosto- mosing bands (Cutler, 1994).
Since thè Cambrian Ottoia prolifica , Walcott 1911 and Louisella pedunculata Walcott, 1911 ffom thè Burgess Shale (British Columbia, Canada) were definitively accepted as pri- apulid worms sharing a common ancestor with sipunculans (Conway Morris, 1977; Cutler, 1994), since Banffia constric- ta Walcott, 1991 ffom thè same outcrop was revised as an arthropod of unknown affinities (Caron, 2001) and since Schizoproboscina ivanovi Yakovlev, 1939 from thè Carboniferous of Russia and Stoma hians Hadding, 1913 ffom thè Ordovician of Sweden were moved to an uncertain order within polychaetes (Sepkoski, 2002), only thè Silurian Lecthaylus gregarius Weller, 1925 ffom Blue Island (Illinois) and thè Jurassic Epitrachys Ehlers, 1 869 ffom thè Solnhofen Plattenkalk actually preserve evidences supporting a sipuncu- lan affinity (Ehlers, 1869 Weller, 1925; Roy & Croneis, 1931). In comparison with R. scutata n. gen. n. sp., Epitrachys rugosus Ehlers, 1 869, Epitrachys granulatus Ehlers, 1 869 and Lecthaylus gregarius lack thè caudal shield but they show a regularly segmented cuticle. Moreover, Epitrachys has a gran¬ ulose cuticle (Ehlers, 1869), not smooth like that of Rondeletia n. gen., Lecthaylus Weller, 1925 has a smooth cuticle but thè relative dimensions of introvert and trunk are very different if compared with those of Rondeletia n. gen. (Weller, 1925; Roy & Croneis, 1931). According to Cutler (1994), thè remaining sipunculan fossil record includes some ichnogenera ffom Cambrian to Miocene, such as Trypanites Màgdeffau, 1982, Zoophycos Massalongo, 1855, Trichichnus Frey, 1970 and Ancorichnus Heinberg, 1974. Moreover, soli- tary corals with sipunculan symbionts are known ffom thè Cretaceous of Texas and Tennessee and ffom thè Miocene of Florida (Cutler, 1994), even if Stolarski et al. (2001) reported thè exsistence of this symbiosis in older outcrops also. Brett & Cottrell (1982) reported thè presence, in thè Devonian Pleurodictyum Goldfuss, 1829 tabulate coral, of overgrown gastropod shells occupied by a secondary resident, probably a sipunculan. Finally, Hàntzschel (1975) treated as sipunculan thè borings found in some Cypricardia Lamarck, 1819 valves ffom thè Pliocene of Italy and described under thè ichnogenus Sabella Linnè, 1767.
Phylum Hemichordata Bateson, 1885 Class Enteropneusta Gegenbaur, 1 870 Family uncertain
Genus Megaderaion Arduini, Pinna & Teruzzi, 1981 Megaderaion callovianum n. sp.
Fig. 8, PI. IL
Diagnosis : elongate wormlike body clearly divided Ho three regions: elongate, rounded and ogival proboscis v prosome), short and thick subrectangular collar (meso- :eì and vermiform tapering trunk (metasome). Trunk covered by a segmented cuticle.
Etymology : referring to thè geological age of thè La Voulte-sur-Rhòne outcrop.
Geological age: Lower Callovian (Middle Jurassic).
Type locality : La Boissine, La Voulte-sur-Rhòne.
Material : 1 specimen. L.P.M.-B. 48352 (holotype).
Description : thè examined specimen is mostly pre- served as a three-dimensional pyritized relief. The total body length is about 5 1 mm and thè maximum width is 5 mm (in thè mesosome). The ogival proboscis is 8 mm long and 2.5 mm wide at thè proximal extremity, grading to 4.5 mm toward thè base, where it is preserved in more relief. The proboscis and thè collar are separated by a deep and 0. 1 mm wide furrow. On thè left side of thè fur- row a 0.1 mm long and 1.5 mm wide peduncle connect- ing thè proboscis to thè collar is evidenti it probably underwent displacement from its originai position in thè middle of thè furrow.
At thè proximal extremity thè subrectangular and 4 mm long collar is wider (5 mm) than thè proboscis, undergoing a slight narrowing distally (4 mm). The late- ral and posterior margins of thè collar are convex, while thè anterior is concave so that it appears slightly folded around thè base of thè proboscis.
The maximum width of thè trunk is at thè collar-trunk boundary, where it measures 5 mm, grading to 1 mm dis¬ tally. The trunk appears divided into two distinct regions separated by a slight narrowing: a subcylindrical and 20 mm long region immediately behind thè collar, that decreases in width toward thè posterior extremity (5-2.5 mm), and an elongate (17 mm), more flattened and nar- rower posterior region (1.5-1 mm). The anterior section of thè trunk is preserved as an imprint in its posterior half and, as in living enteropneusts, presumably consisted of thè branchio-genital, oesophageal and hepatic regions. The boundaries of these regions cannot be distinguished
Fig. 8 - Megaderaion callovianum n. sp., holotype (L.P.M.-B. 48352, x 4).
POLYCHAETE, SIPUNCULAN AND ENTEROPNEUST WORMS FROM THE LOWER CALLOVIAN
13
in thè fossil. The posterior section may represent thè intestinal region. This last appears clearly divided into several thin metameres. The trunk metameres are gradu- ally less evident toward thè collar, behind which they are evident only on thè right side of thè body: this suggests that a segmented cuticle covered thè trunk.
Remarks: thè three distinct regions (proboscis, collar and trunk) that comprise thè body of thè specimen justify its assignment to thè class Enteropneusta within thè phylum Hemichordata, which also includes thè Planctosphaeroidea Spengel, 1932 and Pterobranchia Lankester, 1877. Both thè Planctosphaeroidea and Pterobranchia have recently been suggested to be included in thè Enteropneusta (Harrison & Ruppert, 1997; Cameron et al., 2000). The Enteropneusta are generally subdivided into three families, Spengeliidae Willey, 1899, Ptychoderidae Spengel, 1893 and Harrimannidae Spengel, 1901, on thè basis of thè stomo- chord architecture and other anatomical characters, such as liver sacs and nerve roots, structures that have a very low preservation potential (Cameron, 2002). Only two fossil enteropneusts have been described previously: M. sine- muriense Arduini, Pinna & Teruzzi, 1981 from thè Sinemurian (Upper Jurassic) of Osteno (Lombardy, Italy) and Mazoglossus ramsdellii Bardack, 1997 from thè Carboniferous of Mazon Creek, Illinois. This poor fossil
record is probably due to thè very fragile nature of thè enteropneust body (Cameron, 2002). The enteropneust fos¬ sil record includes some Paleozoic traces referred to thè ichnogenus Nereites MacLeay in Murchinson, 1839 (Mangano et al., 2000). A Paleozoic enteropneust has been discovered in thè Middle Cambrian Burgess Shale (Boulter, 2003), but it is stili undescribed.
The assignment of thè examined material to thè genus Megaderaion is supported by thè shape and relative dimen- sions of thè prosome, mesosome and metasome. These are very different from those of Mazoglossus Bardack, 1997, which has a wider, less elongate proboscis together with a differently shaped collar (Bardack, 1997). A prosome twice as long as thè mesosome, together with a rather elongate proboscis, are very similar in their morphology to those of thè living Harrimania planktophilus Cameron, 2002, sup- porting thè hypothesis (Arduini et al., 1981) that Megaderaion is related to thè Harrimanidae, thè family including thè most primitive species of living enterop¬ neusts. The assignment of thè Megaderaion specimen from La Voulte-sur-Rhòne to a different species is justified main- ly on thè different shape of thè collar. Morover, in contrast to that in M. sinemuriense, thè anterior extremity of thè pro¬ boscis of M. callovianum n. sp. appears not angular but rounded, with a short peduncle at thè base.
UNRELATED SPECIMENS
The specimens L.P.M.-B.48346, L.P.M.-B. 48348, L.P.M.-B.48349, L.P.M.-B.48350 (part and counterpart) and L.P.M.-B. 48351 represent undetermined taxa within thè class Polychaeta. Only for L.P.M.-B. 48348 and L.P.M.-B. 48351 can an assignment within thè order Capitellida be suggested. They are similar to Iubarenicola fìscheri n. gen. n. sp. in thè generai body morphology and preservation, since they too have a subconical prosto- mium and thè body is preserved coiled posteriorly, but it is impossible to prove a capitellid affinity. L.P.M.-B. 48348 measures about 20 mm in length and 3 mm in max¬ imum width in thè centrai body section. L.P.M.-B.48351 . is about 80 mm long and 4 mm wide along its length.
Noteworthy also is thè fact that L.P.M.-B.48346, a 150 mm long and 5 mm wide polychaete largely preserved as an imprint, seems to be preserved with its pharynx everted. The pharynx imprint is about 1 1 mm long and 7 mm wide. The generai body morphology of this specimen, hearing an undetermined number of very elongate chaetae (up to 15 mm) on each metamere, is reminiscent of thè eunicemorph polychaetes but as thè stomodaeal armature is not pre¬ served, it is impossible to confirm its affinity.
L.P.M.-B.48347 (Fig. 9) remains undetermined, but it does not show any polychaete character. It lacks both thè caudal and cephalic extremity and it does not show metamerism or chaetae. The preserved body fragment measures about 70 mm in length and 3 mm in width. The • major features of this worm are thè dorso-ventrally flat- tened body with an elliptical section and thè presence of four thin longitudinal ridges developed along thè preserved body length: these characters are consistent with any group within thè phylum Nemertea. In particular thè thin ridges may represent thè longitudinal muscle bundles that are localized under thè epithelium in thè nemertines.
Fig. 9 - L.P.M.-B.48347, undetermined worm (x 1 .6).
14
ANNA ALESSANDRELLO, GIACOMO BRACCHI & BERNARD RIOU
DISCUSSION AND CONCLUSIONS
The examined material includes 10 polychaetes (2 arenicoloideans and 8 aphroditoideans), 6 sipunculans and 1 enteropneust. Six specimens stili remain undeter- mined: they may be related to polychaetes apart from one specimen which may be a nemertean.
The La Voulte-sur-Rhòne biota is noteworthy for thè early diagenetic mineralization that results in calcite replacement, phosphatization or pyritization (Wilby & Briggs, 1997; Wilby et al., 1995 and 1996; Wilby, 2001) and thus mostly preventing thè collapse or compaction of organic walls (Grimes et al., 2001). Two interpretations of thè La Voulte-sur-Rhòne paleoenviroment (Dietl & Mundlos, 1972; Fischer & Riou, 1982a; Fischer, 2003) have been offered: thè first is a Sargasso sea model where thè benthic fauna inhabited carpets of floating algae, while thè second involves a hypersaline lagoon. The soft tissues of some of thè specimens described here, such as thè lubarenicola fìscheri n. gen. n. sp. holotype and thè Rondeletia scutata n. gen. n. sp. paratype, are preserved as pyrite coats in which thè internai structures are not evident. These alone allow some considerations of thè La Voulte- sur-Rhòne paleoenviroment. According to Allison (1990) and Canfield & Raiswell (1991), these minerai coats formed through thè pyritization by anaerobio sulphate- reducing bacteria that pseudomorph thè soft parts: this process especially occurs in shallow and well-lit enviro- ments, such as estuaries and delta, even if it is not confined to such settings (Briggs et al., 1996; Grimes et al., 2002). An enviroment located in shallow water has been postulat- ed also for thè Devonian Hunsriick Siate, another outcrop known for thè widespread three-dimensional pyritization, even if thè debate about its origin is stili in progress (Etter, 2002b). Moreover, thè presence of enteropneusts and sipun¬ culans, thè body of which is typically very fragile, thè pro- fusion of articulated Bositra buchi valves, thè presence of well-preserved echinoderms, particularly thè ophiuroid Ophiopinna elegans, and thè lack of evidence of current activity, such as ripple marks, clearly suggest a below storni wave base or a stagnant enviroment (Etter, 2002a; Fischer, 2003), such as a restricted lagoon. The abundance of inver- tebrates with articulated hard parts, such as ophiuroids (Dietl & Mundlos, 1972) and crustaceans (Secretan & Riou, 1986; Carriol & Riou, 1991), also indicates that thè preserved organisms of La Voulte-sur-Rhòne probably underwent rapid burial after death, even if this is not a pre¬ requisite for soft tissue pyritization. This kind of fossiliza- tion seems to occur in poorly oxygenated (dysaerobic) bot¬ toni waters rather than in euxinic enviroments, because in dysaerobic waters thè pore fluids circulating in thè sedi- ments contain elevated levels of dissolved iron allowing soft part pyritization (Allison, 1990; Allison & Briggs, 1991; Canfield & Raiswell, 1991; Grimes et al., 2002; Briggs, 2003). According to Dietl & Mundlos (1972), thè rapid burial of thè La Voulte-sur-Rhòne fauna was probably thè result of mudflows, as evidenced by thin individuai lay- ers covering thè bedding levels of articulated invertebrates.
Moreover, thè rapid burial inhibits oxygen diffusion, there- by promoting soft tissue preservation, even if, as previous- ly emphasized, this does not guarantee fossilization. The presence of pelagic and nekto-benthic species at La Voulte- sur-Rhòne supports thè existence of a poorly-oxygenated enviroment, but thè great abundance of benthic taxa indi¬ cates that thè sea bottoni was never anoxic for long. According to Etter (2002a), such paleoecological features reflect periodic sediment settling that resulted in periodic oxygen depletion.
Finally, thè great spread of both epibenthic (such as thè aphroditoideans and reptantian decapods) and endobenthic (such as enteropneusts, sipunculans and arenicoloideans) taxa, thè presence of several nekto-benthic elements (such as rhynchonellid brachiopods, pycnogonids, ophiuroids, mysidacean, penaeid and cumacean crustaceans), thè occurence of relatively few open-water organisms (like actinopterygians, sharks and cephalopods) and thè absence of algae remains, is not consistent, in thè opinion of thè authors, with a Sargasso Sea model (Dietl & Mundlos, 1972; Fischer & Riou, 1982a; Fischer, 2003) for thè La Voulte-sur-Rhòne outcrop.
It is impossible to establish with reasonable certainty whether La Voulte-sur-Rhòne was a deep or a shallow basin. The absence of storm layers seems to indicate that thè sediments settled below storm wave base, while thè high lateral variability of thè Lower Callovian sequence suggests that there was significant relief on thè sea bed (Elmi, 1967; Etter, 2002a). However, in thè opinion of thè authors, this Jurassic Lagerstàtte can be interpreted as a Coastal lagoon, also because of thè presence of some Con¬ tinental elements, such as a crocodilian skull, spores and leaves (Fischer, 2003), thè great abundance of Bositra buchi, which is no longer interpreted as a pseudoplank- tonic bivalve (Etter & Tang, 2002), thè presence of pyriti¬ zation, which can imply shallow water, and thè presence of terrigenous clastics that were deposited in thè epicon- tinental sea of thè La Voulte area following erosion from thè eastern margin of thè Massif Central (Etter, 2002a).
Acknowledgements
W e thank Jean-Claude Fischer (Muséum National d’Histoire Naturelle de Paris, Laboratoire de Paléontologie), who allowed us to study thè examined material, Giorgio Teruzzi (Museo Civico di Storia Naturale di Milano, Department of Paleontology), Derek E.G. Briggs (Yale University, Department of Geology and Geophysics) and Maria Cristina Gambi (Stazione Zoologica ‘Anton Dòhrn’ di Napoli, Benthic Ecology Laboratory) for thè useful advices and careful review of thè manuscript, Fabio Fogliazza and Luciano Spezia (Museo Civico di Storia Naturale di Milano) for drawings and photographs. We also wish to thank Alvaro Invernizzi and thè Gilardoni’s engineers for thè cooperation in thè radiographic investigation.
The opinions expressed in thè paper remain our own.
'I
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Anna Alessandrello & Giacomo Bracchi: Sezione di Paleontologia, Museo Civico di Storia Naturale, Corso Venezia 55, 20121 Milano, Italy.
e-mail: anna.alessandrello@comune.milano.it; mojak@inwind.it t 'Ornard Riou. Musée de Paléontologie de La Voulte, 4 quai Anatole, 07800 La Voulte-sur-Rhòne, France.
_ e-mail: museefossiles@cegetel.net
. sipunculan and enteropneust worms from thè Lower Callovian (Middle Jurassic) of La Voulte-sur-Rhòne (Ardèche, France) M.n oue della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXII - Fascicolo I
*
Hammatoceratidae (excl. Phymatoceratinae ) Hildoceratidae (excl. Hildoceratinae e Bouleiceratinae). pp. 1-70. 2 tavv. n.t.. 6 Jìgg.. 6 tavv.
II - VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi
dell’Anisico superiore di Lenna in Val Brembana (Bergamo)./?/?. 71- 142. 5 Jìgg.. 11 tavv.
III - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell'Alpe Turati (Erba. Como). Generi Hildoceras, Phymatoceras. Paroniceras e Frechiella. Conclusioni generali, pp. 143-204. 2 figg.. 6
tavv.
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Meneghini nelle Tavv. 1-22 della « Monographie desfossiles dii calcai- re muge ammonitique » (1867-1881 ). pp. 5-22. 2 Jìgg.. 6 tavv.
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d'Orta). pp. 23-92. 42 fìgg.. 4 tavv. n.t.
Ili - PETRUCCI F„ BORTOLAMI G. C. & DAL P1AZ G. V., 1970 - Ricerche sull'anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul suo substrato cristallino./?/?. 93-169. con carta a colori al 1:40.000. 14 Jìgg . 4 taw. a colori e 2 b.n.
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I - CANTALUPPI G.. 1970 - Le Hildoceratidae del Lias medio delle regio¬
ni mediterranee - Loro successione e modificazioni nel tempo. Riflessi biostratigrafici e sistematici, pp. 5-46. con 2 tabelle ne! testo.
II - PINNA G. & LEVI-SETTI F.. 1971 - 1 Dactvlioceratidae della
Provincia Mediterranea (Cephalopoda Ammonoideaj. pp. 47-136. 21 figg.. 12 tavv.
Ili - PELOSIO G.. 1973 - Le ammoniti del Trias medio di Asklepieion (Argolide. Grecia) - I. Fauna del «calcare a Ptychites » (Anisico sup.). pp. 137-168. 3 Jìgg.. 9 taw
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Problematica ed ergologia di una facies dell'Eneolitico Padano, pp. 5- 80. 2 figg.. 20 tavv.
II - PETRUCCI F. 1972 -Il bacino del Torrente Cinghio (Prov. Parma).
Studio sulla stabilità dei versanti e conservazione del suolo, pp. 81-127. 37 figg.. 6 carte tematiche.
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(Bergamo)./?/?. 5-34. 16 Jìgg.. 16 tavv.
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d'Arda (Piacenza. Italia)./?/?. 35-78. 6 Jìgg.. 5 tavv. ili - BRAMBILLA G., 1976 - I Molluschi pliocenici di Villalvernia Alessandria). I. Lamellibranchi. pp. 79-128. 4 Jìgg.. 10 tavv.
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- CORNAGGIA CASTIGLIONI O. & CALEGARI G.. 1978 - Corpus delle pintaderas preistoriche italiane. Problematica, schede, iconogra¬ fia. pp. 5-30. 6 figg.. 13 taw.
- PINNA G.. 1979 - Osteologia dello scheletro di Kritosaurus notabilis (Lambe. 1914) del Museo Civico di Storia Naturale di Milano ( Omithischia Hadrosauridaej. pp. 31-56. 3 figg.. 9 taw.
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Volume XXIII
- GIACOBINI G.. CALEGARI G. & PINNA G.. 1982 - I resti umani fos¬
sili della zona di Arena Po (Pavia). Descrizione e problematica di una serie di reperti di probabile età paleolitica, pp. 5-44. 4 Jìgg.. 16 taw.
|l - POLUZZI A.. 1982 - 1 Radiolari quaternari di un ambiente idrotermale del Mar Tirreno, pp. 45-72. 3 Jìgg.. 1 lab.. 13 tavv.
II - ROSSI F.. 1984 - Ammoniti del Kimmeridgiano superiore Berriasiano inferiore del Passo del Furio (Appennino Umbro-Marchigiano)./?/?. 73- 138. 9 Jìgg.. 2 tabb.. 8 taw.
Volume XXIV
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sauro triassico del sottordine Lacertilia. pp. 7-28. 12 Jìgg.. 2 taw.
Il - NOSOTTI S-. PINNA G.. 1989 - Storia delle ricerche e degli studi sui rettili Placodonti. Parte prima 1830-1902./?/?. 29-86. 24 figg.. 12 taw.
Volume XXV
i - CALEGARI G.. 1989 - Le incisioni rupestri di Taouardei (Gao. Mali).
1 Problematica generale e repertorio iconografico./?/?. 1-14. 9 figg.. 24 taw. - PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia funzionale e paleoecologia del rettile placodonte Psephoderma alpinum Meyer. 1858 . pp. 15-50. 20 figg.. 9 ta\s
I - CALDAICA R.. 1990 - Rev isione Tassonomica delle specie paleartiche del genere Tvchius Germar (Coleoptera Curculionidae). pp. 51-218.
575 Jìgg
Volume XXVI
' - PINNA G.. 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia. Placodontia )./?/?. 1-21. 23 figg.
- CALEGARI G. a cura di. 1993 - L'arte e l'ambiente del Sahara preisto¬ rico: dati e interpretazioni./?/?. 25-556. 647 figg.
Ili - ANDR1 E. e ROSSI F., 1993 - Genesi ed evoluzione di frangenti, cin¬ ture. barriere ed atolli. Dalle stromatoliti alle comunità di scogliera moderne, pp. 559-610. 49 Jìgg.. 1 tav.
Volume XXV II
I - PINNA G. & GHISELIN M. edited by, 1996 - Biology as History. N. 1.
Systematic Biology as an Historical Science, pp. 1-133. 68 figg.
II - LEONARDI C. & SASSI D. a cura di, 1997 - Studi geobotanici ed ento-
mofaunistici nel Parco Regionale del Monte Barro, pp. 135-266.
Volume XXV 111
1 - BANFI E. & GALASSO G., 1998 - La flora spontanea della città di Milano alle soglie del terzo millennio e i suoi cambiamenti a partire dal 1700 .pp. 267-388.
Volume XXIX
1 - CALEGARI G.. 1999 - L'arte rupestre dell'Eritrea. Repertorio ragiona¬ to ed esegesi iconografica./?/?. 1-174.
Volume XXX
I - PEZZOTTA F. edited by, 2000 - Mineralogy and petroiogy of shallow depth
pegmatites. Paper from thè First International Workshop, pp. 1-117.
II - PARISI B., FRANCHINO A. & BERTI A. con la collaborazione di
POTENZA B. & RUBINI D.. 2000 - La Società Italiana di Scienze Naturali 1855 - 2000. Percorsi storici, pp. 1-163.
Ili - DE ANGELI A. & GARASSINO A., 2002 - Galatheid, chirostylid and porcellanid decapods (Crustacea, Decapoda, Anomura) from thè Eocene and Oligocene of Vicenza (N ltaly). pp. 1-40.
Volume XXXI
I - NOSOTTI S. & RIEPPEL O., 2002 - The braincase of Placodus Agassiz,
1833 (Reptilia. Placodontia). /?/?. 1-18.
II - MARTORELL1 G.. 2002 - Monografia illustrata degli uccelli di rapina
in Italia. (1895). Riedizione a cura di Fausto Barbagli, pp. [XX] 1- 216. Ili - NOSOTTI S. & RIEPPEL 0., 2003 - Esaurosphargis dalsassoi n.gen.n.sp.. a new, unusual diapsid reptile from thè Middle Triassic of Besano (Lombardy. N ltaly)./?/?. 1-33.
Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali, Museo Civico di Storia naturale, Corso Venezia 55 - 20121 Milano Pubblicazione disponibile al cambio
MEMORIE
della Società Italiana di Scienze Naturali e del Museo Civico
Volume XXXII - Fascicolo II di Storia Naturale di Milano
OLIVER RIEPPEL & JASON J. HEAD
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ,
FROM CENOMANIAN (LATE CRETACEOUS) OF LEBANON
MILANO DICEMBRE 2004
INDEX
Introduction Pag. 3
Abbreviations used in thè figures Pag. 4
Material Pag. 5
The geological provenience of Eupodophis Pag. 5
Description of thè material Pag. 6
Specimen MSNM V 3660 Pag. 6
Specimen MSNM V 3661 Pag. 9
Specimen MSNM V 4014 Pag. 14
Discussion Pag. 20
Systematic conclusions Pag. 22
General conclusions Pag. 24
Acknowledgements Pag. 24
References Pag. 24
© 2004 Società Italiana di Scienze Naturali Museo Civico di Storia Naturale Corso Venezia, 55 - 20121 Milano
In copertina: Eupodophis descouensi. Fotografia di Luciano Spezia.
Registrato al tribunale di Milano al n. 6694
Direttore responsabile: Anna Alessandrello Responsabile di redazione: Stefania Nosotti Grafica editoriale: P.R.G.
Stampa: Litografia Solari, Peschiera Borromeo - Dicembre 2004
ISSN 0376-2726
Olivier Rieppel & Jason J. Head
New specimens of thè fossil snake genus Eupodophis Rage & Escuillié, from Cenomanian (Late Cretaceous) of Lebanon
Abstract - Three new specimens of thè mid-Cretaceous marine snake, Eupodophis descouensi Rage & Escuillié, from Lebanon are described in detail, and thè phylogenetic affinities of this taxon are discussed. Two of thè three specimens represent nearly complete, articulated skeletons of imma¬ ture individuals. Eupodophis shares thè presence of well-developed hind limbs with Pachyrhachis and Haasiophis, both marine mid-Cretaceous snakes from thè Middle East. Eupodophis is unique amongst thè Cenomanian snakes with well-developed hind limbs in having distally fuseci, but proximal- ly bifurcating chevrons in its relatively very short tail. The type of articulation of these chevrons with thè vertebral centra is autapomorphic for Eupodophis, and cannot be compared to varanoid or any other non-ophidian squamate (Tizard’) condition. The skull of Eupodophis most closely resembles that of Pachyrhachis, whereas its vertebral morphology most closely approaches that of Simoliophis and Mesophis. The status of these mid- Cretaceous basai macrostomatans, and their interrelationships are discussed.
Key-words - Reptilia, Lepidosauria, Serpentes, fossil snakes, Cretaceous, Cenomanian, phylogeny.
Riassunto - Nuovi esemplari del serpente fossile Eupodophis Rage & Escuillié (Cenomaniano, Cretacico superiore, Libano).
Vengono descritti in dettaglio tre nuovi esemplari del serpente marino Eupodophis descouensi Rage & Escuillié provenienti dal Libano e vengo¬ no discusse le affinità filogenetiche del taxon. Due dei tre esemplari sono scheletri articolati quasi completi appartenenti a individui immaturi. Eupodophis condivide con Pachyrhachis e Haasiophis, serpenti marini del Cretacico superiore del Medio Oriente, la presenza di arti posteriori ben sviluppati. Caso unico tra i serpenti cenomaniani con arti posteriori ben sviluppati, nella breve coda Eupodophis possiede chevrons fusi distalmente, ma biforcati in prossimità dei corpi vertebrali. La modalità di articolazione di tali chevrons con i centra vertebrali rappresenta un’autapomorfia di Eupodophis, e non può essere comparata alla condizione che si osserva nei varanoidi o in qualsiasi altro squamato non-ofidiano (“lucertole”). Il cra¬ nio di Eupodophis è molto simile a quello di Pachyrachis, mentre le vertebre sono morfologicamente molto simili a quelle di Simoliophis e Mesophis. Vengono infine discussi lo status di questi macrostomati basali del Cretacico superiore, nonché le loro relazioni filogenetiche.
Parole chiave - Reptilia, Lepidosauria, Serpentes, serpenti fossili, Cretacico, Cenomaniano, filogenesi.
INTRODUCTION
The origin of snakes is a longstanding problem in her- petology (Bellairs & Underwood, 1951; Rieppel, 1988), a problem that has recently taken center stage again in thè form of an intensive debate about thè hearing of mid- Cretaceous snakes with well-developed hind limbs on snake origins (for a detailed review of this debate see Rieppel et al., 2003). The genera relevant to this debate are: (i) Pachyrhachis Haas (1979; see also Haas, 1980a, b) from thè Cenomanian of ‘Ein Yabrud (Westbank, Middle East), re-described as thè most basai snake and as sister-group to all other, extant, snakes by Caldwell and Lee (1997; see also Lee & Caldwell, 1998, Caldwell, 2000; Zaher, 1998; Zaher & Rieppel, 2002); (ii) * Eupodophis Rage & Escuillié (2000; see also Rage & Escuillié, 2001, 2003), from thè Cenomanian of Al Namoura (Lebanon), described as a basai snake with a skull anatomy that is convergent on macrostomatans; and l(iii) Haasiophis Tchemov et al. (2000; see also Rieppel et al., 2003) from thè Cenomanian of ‘Ein Yabrud '(Westbank, Middle East), described as a basai macros- tomatan with well-developed hind limbs. All these snake
taxa combine thè presence of well-developed hind limbs with a macrostomatan skull structure (Coates & Ruta, 2000; Greene & Cundall, 2000), a combination of charac- ters that is difficult to reconcile with thè current under- standing of snake phylogeny as based on morphology (Fig. 1). Among extant snakes, thè highly autapomorphic scolecophidians (blind snakes and thread snakes) are con- sidered to be thè sister-group of Alethinophidia; within Alethinophidia, thè (probably paraphyletic) ‘anilioids’ (pipe snakes and shield tails) are thè (possibly successive) sister-group(s) of Macrostomata (Cundall et al., 1993; Cundall & Greene, 2000). Loxocemus, Xenopeltis, and thè ‘booids’ (boas, pythons, and their relatives) constitute basai macrostomatans, whereas colubroids (colubrid snakes, cobras, sea snakes, and vipers) represent derived macrostomatan clades. Most scolecophidians, ‘anilioids,’ and basai macrostomatans retain either some pelvic girdle rudiments and/or hind limb rudiments (Kley et al., 2002), but these elements are not nearly as well-developed as they are in thè three fossil snake genera listed above. By contrast, these fossil snakes show a skull structure resem-
4
OLIVIER RIEPPEL & JASON J. HEAD
Fig. 1 - The phylogenetic interrelationships of extant snakes based on morphology. Anilioidea and ‘Booidea’ are putatively paraphyletic groups. Cundall & Greene (2000) exclude Xenopeltis and Loxocemus from Macrostomata.
bling that of macrostomatan snakes, allowing for a greater degree of jaw kinesis than is characteristic of ‘anilioids’ (Cundall, 1995; jaw kinesis in scolecophidians is highly autapomorphic: Kley, 2001; Kley & Brainerd, 1999). The enigma created by this character combination was aptly summarized by Rage & Escuillié (2000: 515): “If bipedal snakes are ready basai Ophidia, then thè distribution of thè macrostomatan structure of thè skull within Ophidia is problematic.” Note that Ophidia was used by Rage & Escuillié (2000) in thè sense of Lee & Caldwell (1998), who grouped thè bipedal fossil snakes as sister-taxa of extant snakes, referring to thè whole assemblage as Ophidia, and to thè extant snakes - nested within Ophidia - as Serpentes. Given thè morphology based topology of snake phylogeny for extant taxa (Greene & Cundall,
2000; see also Scanlon & Lee, 2000; Lee & Scanlon, 2002), thè skull anatomy of thè fossil snakes with well- developed hind limbs optimizes unequivocally as conver- gent upon thè skull anatomy of basai macrostomatans if these fossil snakes are positioned basai to all extant snakes.
The description of Eupodophis by Rage & Escuillié (2000) introduced two characters that had not previously been discussed in arguments on thè phylogenetic relation- ships of thè fossil snakes with well-developed hind limbs, viz. mediai frontal pillars, and articulated chevrons. All alethinophidian snakes are characterized by mediai down- growths of thè frontals, separating thè two olfactory tracts from one another at thè anterior end of these elements (e.g., Rieppel, 1988). Similar mediai frontal flanges are absent in scolecophidians, and they were (tentatively) claimed to be absent in Eupodophis by Rage & Escuillié (2000). The absence of such mediai frontal flanges would lend important support to a basai position of that taxon relative to alethinophidians. A basai position of Eupodophis relative to all extant snakes was furthermore claimed by Rage & Escuillié (2000) to be supported by thè presence of distally fused, but proximally bifurcated chevrons that articulate (i.e., are not fused) with thè cau- dal vertebrae. Such lizard-like articulated chevrons were also claimed to be present in thè snake genus Wonambi from thè Pleistocene of South Australia by Scanlon & Lee (2000), who referred that taxon to thè Madtsoiidae, a putative family of fossil snakes that they considered basai to all extant snakes. A review of thè skeletal anatomy of Wonambi by Rieppel et al. (2002) confirmed thè “booid” status of thè latter genus (Barrie, 1990), and thè coding of distally fused yet proximally articulating chevrons for both Wonambi and Eupodophis in a new phylogenetic analysis did not cause these two snake genera to group outside basai extant snakes (Rieppel et al., 2002). Instead, they grouped with Pachyrhachis and Haasiophis as basai macrostomatans again.
In this paper we describe thè skeletal anatomy of three additional specimens of Eupodophis from thè middle Cenomanian of Lebanon, kept at thè Museo Civico di Storia Naturale di Milano, and discuss their phylogenetic significance.
ABBREVIATIONS USED IN THE FIGURES
ang |
angular |
f.c |
facet on compound bone for coronoid |
avt.p |
anteroventral process |
fe |
femur |
bo |
basioccipital |
fi |
fibula |
bs |
basisphenoid |
fp.prf |
foot process of prefrontal |
c |
coronoid |
hy |
hypapophysis |
chv |
Chevron |
il |
ilium |
cn |
vertebral centrum |
is |
ischium |
com |
compound bone (of lower jaw) |
izr |
interzygapophyseal ridge |
d |
dentary |
lf.f |
lateral frontal flange |
ec |
ectopterygoid |
ls |
laterosphenoid |
eo |
exoccipital |
m |
maxilla |
f |
frontal |
Me |
MeckeTs canal |
L.
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
5
mf.f |
mediai frontal flange |
so |
supraoccipital |
mp.m |
mediai (palatine) process of maxilla |
sp |
splenial |
mp.pl |
mediai (choanal) process of palatine |
st |
supratemporal |
n |
nasal |
stp |
stapes |
nc |
neural canal |
syn |
synapophysis |
ns |
neural spine |
ti |
tibia |
P |
parietal |
zyg |
zygantrum |
pel |
pedicel for Chevron articulation |
zys |
zygosphene |
pev.f |
pelvic fragments |
V |
vomer |
pi |
palatine |
vl9 |
1 9th precloacal vertebra |
pm |
premaxilla |
v22 |
22nd precloacal vertebra |
pmf |
posterior mylohyoid foramen |
v37 |
37th precloacal vertebra (as preserved) |
po |
postorbitai |
v40 |
40th precloacal vertebra (as preserved) |
poz |
postzygapophysis |
v76 |
76th precloacal vertebra (as preserved) |
prf |
prefrontal |
v78 |
78th precloacal vertebra (as preserved) |
prò |
prootic |
vi 15 |
1 1 5th precloacal vertebra |
prz |
prezygapophysis |
vi 17 |
1 1 7th precloacal vertebra |
Pt |
pterygoid |
vl41 |
1 4 1 st precloacal vertebra |
q |
quadrate |
vl43 |
143rd precloacal vertebra |
rb |
rib |
vl46 |
146th precloacal vertebra |
r.prl |
recess for paralymphatic System |
vl50 |
150th precloacal vertebra |
sc.f |
subcentrai foramen |
vl51 |
1 5 lst precloacal vertebra |
sm |
septomaxilla |
vl55 |
155th precloacal vertebra |
MATERIAL
MSNM V 3660: a series of anterior, middle, and pos- terior precloacal vertebrae, measuring 590 mm in length (620 mm in Dal Sasso & Renesto, 1999; three measure- ments of thè specimen yielded values between 587 mm and 591 mm as measured from thè anterior end of thè first preserved vertebra to thè posterior end of thè posterior precloacal vertebral column as it re-emerges from thè mid-precloacal vertebral column for thè second time).
Middle Cenomanian, Haqel outcrop, Lebanon.
MSNM V 3661: a nearly complete, articulated speci¬ men measuring 330 mm in length (300 mm in Dal Sasso & Renesto, 1999). Middle Cenomanian, Namoura out¬ crop, Lebanon.
MSNM V 4014: a nearly complete, articulated speci¬ men measuring 388 mm in length. Middle Cenomanian, Namoura outcrop, Lebanon.
THE GEOLOGICAL PROVENIENCE OF EUPODOPHIS
The specimen of Eupodophis described by Rage & Escuillié (2000, 2003) carne from thè quarries near thè village of Al Namoura (also written as ‘en Nammoura’, or ‘Nammoura’) located in thè mountains on thè south side of thè valley of Nahr Ibrahim, northern Lebanon (Forey et al., 2003: 230). The same locality yielded two of thè spec- imens described here (MSNM V 3661, MSNM V 4014). The third specimen described here comes from thè famous outcrop near Haqel (also written as ‘Hakel’) in Ouadi Haqel in northern Lebanon (Hiickel, 1970; Hemleben, 1977). The geological and depositional char- acteristics of these Lagerstàtten were last commented upon by Dalla Vecchia et al. (2002), and Forey et al. (2003). Both localities produced fossil land plants, vari- ous invertebrates (including an indisputably marine echi- noid from Namoura: Forey et al., 2003: 231), fishes, Eupodophis, and — in thè case of Namoura - a dolichosaur and a theropod dinosaur (Dalla Vecchia et al., 2002). Fossil fishes, including those that document mass mortality events, are more common at thè Haqel than at thè Namoura locality. According to Hiickel (1970), thè Plattenkalke of Haqel were deposited in small basins
(with diameters of up to 250 meters) that formed at thè outer margin of thè Continental shelf. In contrast, thè exposure of thè Plattenkalk deposits is much more exten- sive at Namoura, there is no evidence of mass mortality events at that locality. The more abundant remains of land plants suggest that Namoura was closer to nearby land than Haqel (Forey et al., 2003: 231), although this was probably “manifest as emergent islands since thè main Continental coastline was several hundred kilometers to thè south” (Forey et al., 2003: 232; approximately 400 kilometers in thè case of thè Haqel locality according to Hiickel, 1970). Forey et al. (2003: 232) reconstructed thè paleoenvironment at Namoura as “a shallow marine, gen- tly sloping platform abutting islands in thè Tethys”. Hiickel (1970) determined thè age of thè fossiliferous Haqel deposits as later Lower Cenomanian, while Forey et al. (2003: 232), based on Dalla Vecchia et al. (2002), conservatively dated thè fossiliferous deposits at Namoura as thè middle portion of thè Middle Cenomanian, mid-Creataceous (Late Cretaceous).
6
OLIVIER RIEPPEL & JASON J. HEAD
Fig. 2 - Specimen MSNM V 3660 (approx. x 1.18). Photo: Luciano Spezia.
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
7
DESCRIPTION OF THE MATERIAL
Specimen MSNM V 3660
(Figs. 2 - 4)
The specimen consists of a series of more than 1 1 1 vertebrae (of which thè last six are partially concealed) that is approximately 590 mm long (Fig. 2). In its size, this specimen matches thè specimen described by Rage & Escuillié (2000). The specimen comprises anterior, and mid- to posterior precloacal vertebrae. Elongated ribs are associated with thè posteriormost preserved vertebrae, indicating that these stili belong to thè precloacal series. There are no preserved elements of thè siculi, of thè cloa- cal, or of thè caudal region, nor are there preserved ele¬ ments of thè pelvis or hind limb. In part, thè vertebrae and ribs split in two when thè part and counterpart slabs were separateci, rendering preservation rather poor. Those ver¬ tebrae that were not split are exposed in either lateral (anteriorly) or ventral view.
The first 13 (preserved) precloacal vertebrae are char- acterized by an elongated, slender, blade-like (i.e., lateral- ly compressed) and posterodorsally slanting neural spine that rises from thè posterior part of thè neural arch (Fig. 3A). This neural spine caused thè vertebrae to tilt during deposition on thè sediment surface, such that thè verte¬ brae are preserved in right lateral view. Their morpholo- gy matches closely thè anterior precloacal (‘cervical’) vertebrae of Pachyophis, which also tilted during thè embedding process because of thè presence of a tali, slen¬ der and blade-like neural spine and hence are exposed in lateral view in thè prepared fossil (Nopcsa, 1923a). The morphology of these vertebrae also closely resembles that of thè “posterior precloacal vertebra” of Simoliophis liby- cus described and figured by Nessov et al. (1998). These latter vertebrae closely match thè morphology of thè ante¬ rior precloacal vertebrae as seen in thè articulated speci- mens of Pachyophis (Nopcsa, 1923a) and Pachyrhachis (Lee & Caldwell, 1998), which confirms their nature as anterior, rather than posterior, precloacal vertebrae. A similar neural spine, thin and slender at its broken base, was inferred to have been present in Simoliophis ( S . rochebrunei sensu Rage, 1984) by Nopcsa (1925).
The anterior precloacal vertebrae rapidly increase in size posteriorly, as was also recognized to be thè case in Pachyophis (Nopcsa, 1923a), Mesophis (Bolkay, 1925), Simoliophis (Nopcsa, 1925), and Pachyrhachis (Lee & Caldwell, 1998). The centrum length of thè well-pre- served eighth element in thè series is 3.3 mm. The ventral margin of thè centrum of these anterior precloacal verte¬ brae is concave in lateral view. Towards its anterior end, thè centrum carries a weakly developed and confluent I dia-parapophysis (synapophysis), which only weakly pro- ■urudes from thè centrum, and which faces in a lateroven- Jtral direction.
As is also thè case in Simoliophis (Nopcsa, 1925), thè anterior precloacal vertebrae carry no distinct hypa- Ipophyses. Nopcsa (1925) described short, ‘knob-like’ [hypapophyses projecting from thè posteroventral margin of thè centrum of larger anterior precloacal vertebrae. A
Fig. 3 - A) Anterior precloacal vertebra (8th preserved element) of spe¬ cimen MSNM V 3660; B) Anterior precloacal vertebra (8th element) of specimen MSNM V 4014.
similarly short and knobby hypapophysis is absent in specimen MSNM V 3660, or only very weakly developed as in thè eighth element preserved in thè series (Fig. 3A). A hypapophysis also appears to be absent on thè anterior precloacal vertebra figured for Simoliophis libycus (Nessov et al., 1998), but in thè text Nessov et al. (1998: 269) describe a “small tubercle-like hypapophysis” for their “posterior trunk” (anterior precloacal) vertebrae. In contrast, a short yet distinct hypapophysis was described for thè anterior precloacal vertebrae of Pachyrhachis (Lee & Caldwell, 1998). In generai, thè hypapophyses of thè pachyostotic fossil snakes, if distinct, are smaller, more posteriorly oriented, and more ovoid in cross-section, than in other snakes. These hypapophyses must not be con- fused with thè knob-like posterior termination of a low midventral ridge at thè posterior margin of thè ventral surface of thè centrum that was described for Mesophis (Bolkay, 1925: 129; “ Knoterì', and fig. 3. See below for thè description of thè same feature in thè posterior pre¬ cloacal vertebrae of specimen MSNM V 3660).
As in Pachyophis (Nopcsa, 1923a) and Simoliophis (Nopcsa, 1925), thè pre- and postzygapophyses are connect- ed by a distinct, laterally projecting keel in thè anterior pre¬ cloacal vertebrae in specimen MSNM V 3660. This keel (Fig. 4) has been previously named ’interzygapophyseal ridge’ by Johnson (1955). The articular surface of thè prezy-
8
OLIVIER RIEPPEL & JASON J. HEAD
gapophysis is mildly inclined dorsomedially, matching thè weakly ventrolaterally facing articular surface of thè postzy- gapophysis. As in Pachyophis (Nopcsa, 1923a), Simoliophis (Nopcsa, 1925), Haasiophis (Rieppel et al, 2003) and Pachyrhachis (Lee & Caldwell, 1998), a prezygapophyseal process is absent. The zygosphene is well exposed at thè base of thè neural arch above thè neural canal, with its later- ally facing articular surface oriented almost vertically. Rage & Escuillié (2000) report an anteriorly notched (concave) zygosphenal lamina (zygosphenal tectum) for Eupodophis, a putatively plesiomorphic character (but see Rieppel et al. , 2002 for a discussion), which cannot be assessed in thè ante- rior precloacal vertebrae of specimen MSNM V 3660 due to their exposure in lateral view.
More posterior vertebrae are all exposed in ventral view (some segments of thè string of vertebrae being split hori- zontally). As in Pachyophis (Nopcsa, 1923a), Simoliophis (Nopcsa, 1925; Nessov et al., 1998) and Pachyrhachis
Fig. 4 - A) Anterior precloacal vertebrae of specimen MSNM V 3660 in ventral view; B) Posterior precloacal vertebrae of specimen MSNM V 3660 in ventral view.
(Lee & Caldwell, 1998), these vertebrae are subject to pachyostosis that is most distinctly developed in thè mid¬ dle of thè precloacal skeleton (Fig. 4). From their expo¬ sure in ventral view thè presence of a relatively low neu¬ ral spine can be inferred. This is because thè presence of a tali neural spine would have caused thè orientation of thè vertebrae in lateral view, as is thè case in thè anterior precloacal elements. The transition from thè anterior pre¬ cloacal vertebrae with a relatively tali neural spine exposed in lateral view, and thè more posterior vertebrae exposed in ventral view occurs relatively abruptly in thè region between thè 21st and 23rd element preserved in thè series. Pachyostosis starts to become apparent in thè region around thè 30th element in thè series, and it is prominent in thè 36th and thè following vertebrae through approximately thè 69,h vertebra, after which it becomes progressively less expressed. The vertebrae posterior to thè 85* element are no longer pachyostotic.
Bolkay (1925) has emphasized that thè vertebrae as exposed in ventral view are always wider than long in Mesophis. In specimen MSNM V 3660, thè first well-pre- served vertebra exposed in ventral view is thè 30* ele¬ ment preserved in thè series. Measurements for it and well-preserved succeeding vertebrae are listed in Table 1 .
Table 1 - Measurements of thè best-preserved vertebrae (exposed in ventral view) of specimen MSNM V 3660. Length of thè centrum is measured as thè maximum length of thè exposed centrum from its anterior to its pos¬ terior margin. Width of thè centrum is again measured as thè maximum width of thè exposed centrum across thè synapophyses.
Well preserved vertebrae exposed in ventral view |
length |
width |
width length |
30th |
5.08 |
5.67 |
1.12 |
35th |
4.93 |
7.28 |
1.48 |
4 1 st |
5.65 |
7.56 |
1.34 |
52nd |
5.15 |
8.49 |
1.65 |
62nd |
5.73 |
7.78 |
1.36 |
66th |
5.18 |
7.65 |
1.48 |
70th |
5.22 |
7.05 |
1.35 |
75th |
5.52 |
-6.33 |
1.15 |
79th |
5.24 |
6.21 |
1.19 |
85th |
4.84 |
6.18 |
1.28 |
93rd |
4.77 |
4.85 |
1.02 |
Given thè articulated nature of thè material, and its sometimes incomplete exposure, these measurements must be considered somewhat approximate. They never- theless confirm that thè vertebral centra are wider than long in Eupodophis, and that thè relation of width to length increases with an increasing degree of pachyosto¬ sis in thè mid-precloacal region. The same appears to obtain for Pachyophis with thè vertebrae exposed in dor- sal view (Lee et al., 1999, fig. 1).
Pachyostosis affects not only thè centrum, but also thè parapophyseal part of thè synapophysis, which is located on thè lateroventral aspect of thè anterior end of thè cen-
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
9
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trum in specimen MSNM V 3660. In thè pachyostotic ver- tebrae of Haasiophis , thè smaller diapophyseal part of thè synapophysis is separateci from thè hypertrophied para- pophyseal part by a distinct groove (Rieppel et al., 2003). MSNM V 3660 is not preserved well enough to allow thè assessment as to whether there is a distinct groove that separates thè hypertrophied parapophyseal from a smaller diapophyseal component of thè synapophysis (Fig. 4). The difference in size between thè parapophyseal and diapophyseal components of thè synapophysis is no greater in Pachyrhachis (Lee & Caldwell, 1998) than in thè majority of alethinophidians (Hoffstetter & Gasc, 1969). A groove separating thè parapophyseal and diapophyseal components of thè synapophysis is absent in Pachyophis and in thè Egyptian material of Simoliophis (Nopcsa, 1925), unlike in Simoliophis rochebrunei (Buffrénil & Rage, 1993; Rage, 1984).
The pachyostotic vertebrae, exposed in ventral view show what Nopcsa (1925: 21) described as a “longitudi- nal tripartition of thè ventral surface of thè centrum . . . that characterizes thè anterior pachyostotic vertebrae of thè Egyptian species ...” of Simoliophis. The same mor- phology is observed in Mesophis (Bolkay, 1925, fig. 3). In fact, specimen MSNM V 3660 shows that this triparti¬ tion can be recognized to a variable degree throughout thè series of pachyostotic mid-precloacal vertebrae (Fig. 4). The tripartition results from a pair of shallow longitudinal grooves that demarcate a relatively broad, shallow and blunt medioventral (hemal) ‘keel’ (rather: ridge) from thè collateral parts of thè fiat ventral surface of thè centrum. Within those shallow longitudinal grooves (Fig. 4A), or along their lateral margins (Fig. 4B), lie thè subcentrai foramina which, as in Simoliophis (Nopcsa, 1925), are irregularly shaped and distributed, i.e. not strictly bilater- ally symmetrically arranged. The posterior end of thè midventral keel expands into a rounded, knob-like struc- ture (“ KnoterT of Bolkay, 1925) at thè posterior margin of thè ventral surface of thè centrum, a feature that is most distinctly expressed behind thè 70th vertebra preserved in thè series in specimen MSNM V 3660 (Fig. 4B). The shallow grooves of Nopcsa’s (1925) ‘tripartition’ are recesses for thè paralymphatic System that are present in all alethinophidians and some scolecophidians (Head, personal observation). Similar grooves are also present in Pachyrhachis and Haasiophis, although thè generai appearance of thè subcentrai grooves in MSNM V 3660 is more similar to Mesophis and Simoliophis than to thè pre- vious taxa.
Numerous ribs are preserved, some of them (most notably thè left rib associated with thè 42nd vertebra pre¬ served in thè series) switched so as to expose thè pos- terodorsally directed and rather weakly developed tuber- culum. A similar rib morphology was described for Simoliophis by Nopcsa (1925). As in thè latter genus, as well as in Pachyrhachis (Lee & Caldwell, 1998), thè ribs are curved in their proximal ‘shoulder’ region, but straight distally (see also Rage, 1984). Although some- what thickened in thè proximal shoulder region within thè pachyostotic precloacal section, thè ribs are subject to a lesser degree of pachyostosis in Eupodophis than is char-
acteristic of Pachyophis (Nopcsa, 1923a; Lee et al., 1999), Pachyrhachis (Lee & Caldwell, 1998), or Haasiophis (Rieppel et al., 2003). In that respect, Eupodophis more closely resembles Mesophis (Bolkay, 1925). Nopcsa (1925) mentions distinct pachyostosis proximally in some ribs of Simoliophis.
Bolkay (1925: 130) noticed a marked difference in thè length of thè longest ribs in Pachyophis and Mesophis. He lists thè average length for thè longest ribs as 23.2 -25.5 mm for Mesophis, but ~40 mm for Pachyophis. But to judge from Nopcsa’s (1923a) piate VII (assumed to ren¬ der thè specimen at approximately naturai size), thè longest ribs do not seem to exceed 26 mm in length. The slightly longer ribs of Pachyophis may well reflect thè somewhat larger overall size of thè latter specimen as compared to Mesophis. Almost none of thè ribs are fully exposed in specimen MSNM V 3660, and their distai part is often represented only by an impression in thè matrix, which creates some inexactness of measurements. However, both thè proximal and distai parts of thè left rib associated with thè 81st vertebra (as preserved) are pres¬ ent, connected with each other by an impression in thè matrix that represents thè missing middle section of that rib. The length of that rib is 27.8 mm, which is just a bit longer than thè longest ribs in Pachyophis. This may reflect thè fact thè specimen MSNM V 3660 is of slight¬ ly larger overall size than Pachyophis.
In summary, and as noted by Rage & Escuillié (2000), thè vertebrae of Eupodophis are very closely similar to those of Simoliophis, differing from thè latter only in thè presence of anterior protuberances on thè dorsal surface of thè neural arch, a character which is not exposed in specimen MSNM V 3660 (but see below for thè descrip- tion of specimen MSNM V 4014). The anterior vertebrae of Eupodophis share with those of Pachyophis (Nopcsa, 1923a) and Pachyrhachis (Lee & Caldwell, 1998) thè tali neural spine, and thè ventrally exposed vertebrae of spec¬ imen MSNM V 3660 closely match those of Mesophis (Bolkay, 1925) in details of surface structure and in thè generai tendency of being wider than long.
Specimen MSNM V 3661
(Figs. 5 - 8)
This is a nearly complete, articulated specimen (Fig. 5) that is much smaller than either thè specimen described by Rage and Escuillié (2000), or MSNM V 3660. The total length of MSNM V 3661 measures 314 mm from thè anterior tip of thè skull to thè posterior tip of thè pre¬ served portion of thè tail. The skull measures 13.2 mm from thè posterior tip of thè left supratemporal to thè anterior tip of thè left maxilla, and 12.3 mm from thè pos¬ terior end of thè basicranium (as preserved) to thè anteri¬ or tip of thè left maxilla.
The skull is exposed in ventral view (Fig. 6). Preservation of thè skull is incomplete due to thè separa- tion of thè part and counterpart slabs, and some of thè ele- ments (right dentary, right maxilla) are vaguely indicated as naturai molds only. However, some of thè elements are
10
OLIVIER RIEPPEL & JASON J. HEAD
Fig. 5 - Specimen MSNM V 3661 (approx. x 1.36). Photo: Luciano Spezia
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
well preserved and offer important anatomical detail. The left maxilla is well preserved except for its posteriormost tip, and it shows a total of 1 8 preserved tooth positions; accounting for thè poorly preserved posterior end, thè entire maxilla may nave carried as many as 21 teeth. The anterior end of thè maxilla is weakly curved medially, and its anterior tip is smooth and blunt, indicating thè absence of a suturai contact with thè premaxilla. No trace of a pre- maxilla can be identified, but thè anterior tip of thè left maxilla approaches thè anterior end of thè naturai mold of thè right maxilla closely enough to allow thè inference that thè premaxilla must have been small and narrow, as is also thè case in Pachyrhachis (Lee & Caldwell, 1998) and Haasiophis (Rieppel et al., 2003). As in macrostom- atan snakes such as Python, thè maxilla carries a broad- based and shelf-like mediai (palatine) process, which is broken due to thè underlying (morphologically: overly- ing) apex of thè elongate prefrontal (Frazzetta, 1966, Fig. 1 6). Only a tooth-bearing fragment of thè posterior part of thè right maxilla is preserved.
Of thè left prefrontal, only thè apex that has been pushed through thè mediai (palatine) process of thè max¬ illa is preserved. Crushed bone behind thè mediai (pala¬ tine) process of thè left maxilla, and between thè latter and thè left frontal, may also represent thè prefrontal. On thè right side of thè skull, thè prefrontal is also crushed, but its outlines and its contact with thè anterior margin of
: ig. 6 - The skull of specimen MSNM V 3661 as preserved in ventral iew.
thè frontal are preserved (Fig. 6). The prefrontal is elon¬ gate and slender, more closely resembling that of thè anil- ioid Cylindrophis than that of Python. The posterolateral corner of thè prefrontal appears to be somewhat thick- ened, forming an articular knob (Frazzetta, 1966, fig. 18), which appears to be received by a shallow facet on thè anterolateral edge of thè frontal. The identification of this articulation is somewhat tenuous, as it depends on a vari- able angle of illumination of thè specimen, but if indeed present, it would indicate that thè skull of Eupodophis might have been prokinetic.
Both frontals are preserved in ventral view, although in a crushed condition. They have been forced apart during thè fossilization process. Two features of thè frontal are observable and important. First, both frontals are of equal length, and their length relative to thè rest of thè skull is unusually elongate compared to extant snakes such as ‘anilioids’ or basai macrostomatans. Indeed, due to their shape and proportions, thè frontals are more similar to those of non-ophidian squamates than to those of extant snakes. Such frontals are so far known to occur only in specimen MSNM V 3661 and in Pachyrhachis (Lee & Caldwell, 1998) among snakes. Secondly, thè left frontal shows what appear to be remnants of a crushed anterior mediai pillar that is characteristic of alethinophidian snakes (Fig. 6). Although preservation is not entirely sat- isfactory, it seems that a gap might have persisted between thè mediai and lateral frontal flanges, as is thè case in ‘anilioids’ (Rieppel, 1978); in macrostomatans, thè medi¬ ai and lateral frontal flanges establish contact medioven- tral to thè olfactory tract (Frazzetta, 1966). It should be noted, however, that thè ventral margins of both thè later¬ al and mediai frontal flanges are subject to breakage, such that thè exact nature of their configuration cannot be established.
At thè back end of thè left maxilla, and dose to thè posterior tip of thè left ectopterygoid, thè left postorbitai is exposed (Fig. 6). Its ventral tip is incomplete. The dor- sal head of thè postorbitai is expanded at its articulation with thè lateral wing of thè parietal (which itself is not preserved) in a manner closely comparable to thè postor¬ bitai in Pachyrhachis (Rieppel & Zaher, 2000, fig. 15B). The right postorbitai is also preserved, but less complete- ly so than thè left one. Given thè similarity of thè left pos- torbital with that of Pachyrhachis, a nearly complete pos- torbital margin can be reconstructed for Eupodophis, a macrostomatan feature.
Behind thè frontals, transversely oriented and vertical- ly positioned shards of bone project from thè sediment (Fig. 6). It is difficult, if not impossible, to unequivocally identify these bone fragments, but they may belong to thè parietal. The only but rather unlikely other alternative would be to interpret these bone fragments as parts of thè basipterygoid processes, which would then be ventrally projecting and transversely oriented structures resem¬ bling thè basipterygoid processes in Python.
The supratemporal in Eupodophis projects from below (morphologically: from above) thè preserved basi- cranium on both sides of thè skull, forming a free-ending posterior process for thè suspension of thè quadrate (Fig.
12
OLIVIER RIEPPEL & JASON J. HEAD
6). The left supratemporal is better preserved than thè right, and it shows a slight expansion of thè posterior end of thè free-ending process. A similar expansion of thè posterior end of thè supratemporal is present in Haasiophis (Rieppel et al., 2003), where it is more dis- tinctly developed than in Eupodophis.
Both quadrates are preserved, splayed outwards due to thè dorsoventral compression of thè skull during fos- silization. As preserved, thè quadrates slant slightly anteroventrally, an orientation that is also observed in Haasiophis with a dorsoventral ly crushed skull, and that is attributed to postmortem dislocation. Given thè overall proportions of thè skull (see below for a discussion), thè quadrate was positioned vertically, or slanted slightly pos- teroventrally, as is characteristic of basai alethinophidi- ans, including basai macrostomatans. The dorsal ends of thè quadrates are broadly expanded, but thè shaft of thè quadarates narrows progressively towards thè mandibular condyles (Fig. 6). In having what appear to be a ventrally slender quadrate, specimen MSNM V 3661 resembles Haasiophis more closely than Pachyrhachis , and it also resembles thè specimen of Eupodophis described by Rage & Escuillié (2000). However, another specimen (MSNM
V 4014) described below exhibits a broad quadrate, such that thè question arises whether thè quadrates of MSNM
V 3661 are incompletely ossified. The anterior and poste¬ rior margins of both quadrates appear to be complete in specimen MSNM V 3661 . The posterior margin of thè left quadrate in particular is perfectly preserved and docu- ments thè absence of a stylohyal in MSNM V 3661.
Little morphological detail can be offered for thè der- mal palate. Bone fragments mediai to thè anterior part of thè left maxilla may represent parts of thè palatine and vomer respectively. Nothing is preserved of thè left ptery¬ goid, whereas a tooth-bearing fragment of thè palatine ramus of thè right pterygoid is positioned alongside thè lateral margin of thè right frontal. What might be thè pos¬ terior end of thè quadrate ramus of thè right pterygoid is located alongside thè basicranium. As preserved, it termi- nates in a blunt tip just in front of thè dorsal end of thè quadrate. The problem with this interpretation is that thè quadrate ramus of thè pterygoid would be much more massively developed in specimen MSNM V 3661 than in specimen MSNM V 4014. The questionable strip of bone might therefore also be interpreted as parts of thè crushed basicranium that were originally facing laterally (thè dat¬ erai wings’ of thè para-basisphenoid and adjacent areas). Both ectopterygoids are preserved, thè left one more completely so than thè right one. The ectopterygoid of specimen MSNM V 3661 closely resembles that of Pachyrhachis (Rieppel & Zaher, 2000, fig. 15B). It is a horizontally positioned lamina of bone that slightly expands towards its anterior end (Fig. 6). The anterior end is not exposed as it underlaps (morphologically: overlaps) thè posterior end of thè maxilla. The somewhat narrower posterior end has a complete, rounded margin, and under- lies thè ventral tip of thè postorbital. The complete preser- vation of thè ectopterygoid in specimen MSNM V 3661 refutes thè claim made by Rieppel & Zaher (2000) that thè ectopterygoid is broken across thè posterior end of thè
maxilla in Pachyrhachis. This claim was made to account for thè shortness of thè ectopterygoid, and for its position almost entirely in front of thè ventral tip of thè postor¬ bital. These are characteristics which specimen MSNM V 3661 shares with Pachyrhachis, and which led to thè identification of thè ectopterygoid as a jugal in Pachyrhachis (Lee & Caldwell, 1998; Lee & Scanlon, 2002). However, as will be discussed in more detail below, thè position of thè ectopterygoid almost entirely in front of thè postorbital results from thè unusually elon- gate frontals that carry thè postorbital backwards.
Of thè braincase, only thè crushed ventral surface of thè posterior part of thè basicranium is preserved (other than possible fragments of basipterygoid processes: see thè dis¬ cussion of thè parietal above). However, thè ventral surface of thè basioccipital is macrostomatan in nature (Tchemov et al., 2000), as it shows distinct ventrally projecting crests (Fig. 6). A longitudinal crest served as thè site of origin of thè protractor pterygoidei (Frazzetta, 1966, fig. 30), while laterally positioned and posteriorly concave crests delin¬ eate facets on thè ventral surface of thè basioccipital for thè insertion of hypaxial neck muscles.
Posterior parts of both mandibular rami are preserved on either side of thè skull. The anterior end of thè right dentary is preserved as a naturai mold only; no tooth counts are therefore available for thè lower jaw. The right mandibular ramus shows thè well-preserved coronoid process formed by thè coronoid (crushed in thè left mandibular ramus). The coronoid of specimen MSNM V 3661 is closely similar to that of Pachyrhachis (Lee & Caldwell, 1998), and very different from that of Haasiophis and all extant snakes, in that it forms a broad, high coronoid process of almost rectangular outlines (Fig. 6). Different from Pachyrhachis, thè dorsal margin of thè broad coronoid process slants posteroventrally in speci¬ men MSNM V 3661 . The coronoid is weakly expanded at its base in an anteroventral direction, and it is applied to thè mediai surface of thè compound bone (i.e., not brac- ing thè dorsal aspect of thè surangular as it does in non- ophidian squamatesi Rieppel & Zaher, 2000). The com¬ pound bone is broken on both sides, but preserved in articulation with thè quadrate. The right mandibular ramus preserves thè rounded and “stubby” retroarticular process that is characteristic of alethinophidian snakes (Rieppel & Zaher, 2000).
The teeth in specimen MSNM V 3661 are slender, needle-shaped and recurved, much like small replicas of thè teeth seen in boas and pythons. The small size of thè teeth makes it difficult to identify anterior, posterior, or lateral (as in Pachyrhachis : Lee & Caldwell, 1998) cut¬ ting edges, but in generai thè enamel surface is smooth and lacks thè ridges or fluting seen in Haasiophis (Rieppel et al., 2003).
A total of 163 vertebrae are preserved in specimen MSNM V 3661. Of those, thè first 41 vertebrae are char- acterized by a tali, slender and blade-like neural spine that is located towards thè posterior end of thè neural arch and that projects in a posterodorsal direction (Fig. 7A; no neu¬ ral spine is developed on thè atlantal neural arch). The height of thè neural spine diminishes gradually in thè last
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
Fig. 7 - A) Anterior precloacal vertebrae of specimen MSNM V 3661, as preserved in lateral view; B) Anterior precloacal vertebrae of speci¬ men MSNM V 4014, as preserved in lateral view.
few elements of that series. As in Pachyophis (Nopcsa, 1923a), Pachyrhachis (Lee & Caldwell, 1998), and spec¬ imen MSNM V 3660, thè neural spines have caused thè vertebrae to tilt relative to thè bedding piane, so that they are exposed in lateral view. Separation of thè part and counterpart slabs caused a sagittal splitting of thè verte- bral centra, but not of thè blade-like neural spines. The sagittally split vertebrae expose thè right half of thè zygosphene in mediai view underlapped (as preserved and exposed) by thè zygantrum of thè preceding neural arch (Fig. 7A). The zygosphene is prominently developed in Eupodophis, as was also noted to be thè case in ■ Pachyophis by Nopcsa (1923a). As in specimen MSNM V 3660, there is no indication of thè presence of hypapophy- ses in thè anterior precloacal region, not even as a slight posteroventral protuberance of thè (split) vertebral centra as it is known from Simoliophis (Nopcsa, 1925; but see thè description of specimen MSNM V 4014 below).
Vertebrae posterior to thè first 4 1 elements continue to be embedded with their lateral side parallel to thè bedding piane. Again, thè vertebrae are sagittally split, exposing thè spongy bone inside thè vertebral centrum. The neural canal is relatively very wide (with a vertical diameter that exceeds thè height of thè centrum; see also Fig. 7A), and thè well-developed zygosphenes continue to be exposed in mediai view in these sagittally split vertebrae. There is ) no indication of even low neural spines, which suggests that thè better part of thè vertebrae adhered to thè coun¬ terpart slab (not available). A small neural spine is appar- ent again in thè tail region. It starts to become noticeable on thè 1 54th vertebra, and increases in height towards thè mid-section of thè tail. It is of a peculiar morphology, in that it is a curved structure (concave anteriorly, convex
posteriorly) that rises from thè posterior end of thè neural arch and terminates dorsally in a straight, horizontally oriented margin (see Fig. 12 showing thè same morphol¬ ogy for specimen MSNM V 4014).
The ribs in specimen MSNM V 3661 are not subject to pachyostosis. This may be related to thè small size of thè specimen (see thè discussion of pachyostosis as relat¬ ed to ontogeny in Lee et al., 1999). Along thè entire pre¬ cloacal body, thè ribs are curved in their proximal ‘shoul- der’ region, but straight for most of their length, and they show a slight distai expansion. Rib morphology thus sug¬ gests a lateral compression of thè body (Rage & Escuillié, 2000). As far as can be ascertained, there is no indication of a well-developed tuberculum, as is present in Simoliophis (Nopcsa, 1925), and there is no evidence that ribs were developed beyond thè 1 42nd vertebra.
In specimen MSNM V 3661, thè longest ribs (occur- ring between thè 80,h and 100,h vertebrae) are 16 mm long. As in Pachyrhachis, thè ribs remain relatively long up to a level shortly in front of thè femur. The last distinctly elongated rib is associated with thè 140th vertebra. Its length, as indicated by an impression left in thè matrix, is 6.1 mm, which equals thè length of thè 5 following verte¬ brae (140* to 145*). The ribs associated with thè 1 4 lst and 142nd vertebrae are poorly preserved (poorly ossified?), but they are abruptly much shorter. No ribs are unequivo- cally identifiable beyond thè 142nd vertebra. Next to thè ventral edge of thè 142nd vertebra lie two small splints of bone that appear to represent either 2 short ribs lying par¬ allel to one another, or a broken bifurcating element. Preservation is poor and structural detail difficult to
Fig. 8 - A) The cloacal region of thè vertebral column and thè hind limb as preserved in specimen MSNM V 3661; B) The pelvic rudiments and hind limb as preserved in specimen MSNM V 4014.
14
OLIVIER RIEPPEL & JASON J. HEAD
ascertain (Fig. 8A), but given thè absence of lym- phapophyses behind thè 142nd vertebra (lymphapophyses are also absent in specimen MSNM V 4014), thè struc- tures mentioned above are interpreted as ribs. This inter- pretation leaves Eupodophìs without lymphapophyses (note that thè putative ‘sacrai rib’ of Pachyrhachis is thè first [articulated] lymphapophysis: Lee, 1998). As pre- served in specimen MSNM V 3661, thè dorsal end of thè ilium lies at thè sanie level as thè non-bifurcating short rib associated with thè 1 4 1 st vertebra, i.e., behind thè seem- ingly bifurcating structure that is associated with thè 142nd vertebra. It is also impossible to identify any element in specimen MSNM V 3661 as a putative sacrai vertebra (see Lee & Caldwell [1998] for thè identification of a sacrai vertebra in Pachyrhachis , and thè comments in Rieppel & Zaher [2000] and Rieppel et al. [2003] on this issue), which argues for thè use of ‘precloacal’ rather than ‘presacral’ vertebrae in thè description of Eupodophis.
In thè absence of lymphapophyses it is difficult to demarcate precloacal from cloacal vertebrae. Addi- tionally, Eupodophis appears to lack caudal pleurapophy- ses, a potential autapomorphy of thè taxon. However, Rage & Escuillié (2000) noted thè relatively very short tail of Eupodophis , as well as thè presence of distally fused, but proximally bifurcated chevrons. They consid- ered these an important plesiomorphy of Eupodophis (Rage & Escuillié, 2000: 518). In specimen MSNM V 3661, thè 146,h through 149th vertebrae are peculiar in that they develop a distinct, ventrally projecting sagittal keel on thè ventral surface of thè centrum (Fig. 8A). At thè anterior end of thè centrum, this keel is developed into a medioventrally located anteroventrally projecting, curved (convex anteriorly, concave posteriorly) and blade-like process that terminates in a blunt (not pointed) tip. On thè 150th and 1 5 1 st vertebrae, this same process is reduced to a small but distinct (unpaired) blade-like pedicel (Figs. 8, 12) located medioventrally at thè anteroventral margin of thè centrum. It is with these pedicels that thè first two chevrons articulate, each ‘embracing’ thè pedicel of its associated vertebra with thè proximally bifurcating ‘arms’ (Fig. 8A). In specimen MSNM V 3661, these two anteriormost chevrons are much shorter than more poste- rior ones. In fact, these two anteriormost chevrons form little more than a minute V-shaped structure with little distai extension. Although frequently broken, thè length of thè chevrons that articulate with thè 152nd and more posterior vertebrae is indicated by naturai molds. The chevrons form distinctly elongate, flattened, blade-like structures with a relatively short, proximally bifurcating part. The chevrons slope posteroventrally relative to thè long axis of thè vertebral column. At thè posterior end of thè preserved part of thè tail, thè chevrons are only repre- sented by weak impressions in thè matrix.
Two small rod-shaped elements are apparent next to thè femur (one as a distinct naturai mold only), which are interpreted as remnants of thè rudimentary pelvis (Fig. 8A). The anterior one of these elements is tentatively identified as thè ilium (represented by a naturai mold), thè other as ischium (preserved in bone). The ilium shows a slight basai expansion.
Of thè hind limbs, only one femur is (partially) pre¬ served on thè right side of thè 144"’ to 1 46th vertebral ele¬ ments. The posterior part of thè femur is represented by a distinct naturai mold. The length of thè femur is 2.98 mm (average of three measurements), and it equals almost exactly thè length of thè three consecutive vertebrae lying next to it (as measured through their split centra). The femur is a straight, slender and essentially featureless ele¬ ment with a slightly constricted diaphysis and slightly expanded proximal and distai ends. No other parts of thè limb are preserved.
Specimen MSNM V 4014
(Figs. 3, 7 - 12)
The total length of thè complete and articulated spec¬ imen (Fig. 9) is approximately 388 mm (thè very tip of thè tail is concealed by a loop of thè body). The specimen therefore matches MSNM V 3661 in size, both being much smaller in size than MSNM V 3660 and thè speci¬ men described by Rage & Escuillié (2000). The head length is best assessed by thè length of thè well-preserved right mandibular ramus, which measures 14.9 mm from its symphyseal tip to thè posterior tip of thè retroarticular process.
The skull is again exposed in ventral view, but with only a very limited assortment of bones preserved (Fig. 10). At thè anterior end of thè skull is located thè small and narrow premaxilla with what appears to be a single, medially located (and presumably internally subdivided) premaxillary channel, and short and pointed posterior vomerine processes. The premaxilla of specimen MSNM V 4014 is edentulous, and in its size and proportions closely resembles thè edentulous premaxilla of Haasiophis (Rieppel et al, 2003; contra Tchernov et al., 2000). The premaxilla is not known for Pachyrhachis (Lee & Caldwell, 1998), Pachyophis (Nopcsa, 1923a) or Mesophis (Bolkay, 1925).
Along thè midiine of thè skull, in front of thè right palatine (Fig. 10), paired crests of bone represent thè ven¬ tral exposure of thè medially descending flanges of thè nasals. At a deeper (morphologically more dorsal) level, thè left nasal preserves a triangular part of its dorsal lam¬ ina. Not enough bone is preserved however, to allow thè assessment of thè outlines of thè entire dorsal lamina of thè nasal.
Between thè left nasal and thè anterior tip of thè left mandibular ramus, thè septomaxillary - vomer complex housing Jacobson’s organ is exposed in articulation in ventral view. The septomaxilla protrudes narrowly from below (morphologically: above) thè anterior dentigerous process of thè left palatine (itself pressed against thè left half of thè mandible), but it can be identified clearly as it forms thè lateral margin of thè opening of Jacobson’s organ. The left vomer is almost completely preserved, and it forms thè mediai margin of thè opening of Jacobson’s organ, meeting thè septomaxilla in a straight longitudinal suture in front of and behind that opening. The slender and elongate vomer has a rounded anterior end, a slightly
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCU1LLIE, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
Fig. 9 - Specimen MSNM V 4014 (approx. x 2.28). Photo: Luciano Spezia.
16
OLIVIER RIEPPEL & JASON J. HEAD
damaged lateral margin, and a tapering posterior part that is broken off from thè main body of thè vomer and that ends in a pointed posterior tip.
The anterior dentigerous process of thè palatine, an alethinophidian feature, is preserved on both sides of thè skull, although thè left one is only poorly exposed, being pressed against thè left mandibular ramus (Fig. 10). The right palatine is almost completely preserved, although broken in its anterior part. The anterior dentigerous process carries eight tooth positions. The posterior end of thè palatine forms a notch that engaged in a tongue-and- groove articulation with thè anterior end of thè palatine ramus of thè pterygoid in a configuration that recalls thè ‘booid’ condition. Also resembling thè macrostomatan condition is thè mediai (choanal) process of thè palatine that is narrow and located towards thè posterior end of thè palatine (Fig. 10). It tapers medially as it provides a pas- sage for thè choanal tube, ending in a blunt tip. There was no well-defined contact between thè choanal process of thè palatine and thè vomer in specimen MSNM V 4014.
The quadrate ramus of thè right pterygoid is preserved and well exposed (Fig. 10). The bone shows a weak later¬ al expansion just in front of a laterally facing facet that would have received thè posterior end of thè ectoptery- goid. More anteriorly, thè teeth of thè palatine ramus of thè pterygoid are aligned along a strip of bone that repre- sents thè incompletely preserved anterior part of thè
Fig. 10 - The skull of specimen MSNM V 4014 as preserved in ventral view.
pterygoid, which itself lies alongside thè basisphenoid. Superficial inspection might create thè impression that thè pterygoid in specimen MSNM V 4014 carried a mas- sively developed, laterally projecting transverse process, as indicated by bilaterally symmetrically arranged globu- lar fragments of bone, thè right one situated next to thè weak lateral expansion of thè pterygoid. These fragments of bone represent parts of thè prootic, however, as will be discussed in more detail below.
The posterior end of thè quadrate ramus of thè right pterygoid overlaps thè internai surface of thè broad quadrate. The right quadrate is disarticulated and rotated posterolaterally, which renders thè assessment of its ori- entation and originai shape difficult. It also appears that thè broken distai part of thè shaft of thè stapes adheres to its (topological) anterior margin (Fig. 10). But whatever thè details of thè structure of thè quadrate, specimen MSNM V 4014 resembles Pachyrhachis more closely than MSNM V 3661 or thè specimen described by Rage & Escuillié (2000) in having an unusually broad quadrate. Only thè ventral half of thè right quadrate is preserved, but again it is broader than is seen in specimen MSNM V 3661. Its orientation indicates that thè quadrate was posi- tioned vertically, or slanted slightly posteroventrally. In front of thè left quadrate, thè posterior tips of two rod- shaped elements are exposed. The lateral one of these is relatively broad anteriorly, but tapering to a blunt tip pos- teriorly. It represents thè posterior part of thè quadrate ramus of thè pterygoid. The mediai element is somewhat more delicate, and represents thè distai end of thè shaft of thè stapes.
Of thè basicranium, part of thè crested ventral surface of thè basisphenoid rostrum is exposed, behind which thè ventral surface of another disarticulated and rotated part of thè basicranium (?) emerges (Fig. 10). The cresting on thè ventral surface of thè basisphenoid is again a macros¬ tomatan feature of specimen MSNM V 4014. The part of thè basisphenoid that is preserved indicates thè presence of a broad, anteriorly tapering para-basisphenoidal ros¬ trum in MSNM V 4014. Little detail can be offered with respect to more posterior parts of thè basicranium, except that a weakly demarcated knob-like projection is present at what presumably constitutes its posterior margin. This projection is too small to represent thè occipital condyle located at thè posterior margin of thè basioccipital. In Haasiophis (Rieppel et al., 2003), a knob-like projection marks out thè posterior end of thè midventral longitudinal crest on thè basicranium (basioccipital). In specimen MSNM V 4014, there is no distinct crest, however, that leads up to this small posterior knob-like projection.
On thè other hand, a small foramen is seen to pierce thè element which, if interpreted as a foramen for a root of thè hypoglossal nerve, would identify thè element as part of thè exoccipital. On that interpretation, thè hypoglossal foramen would be located slightly in front of and above thè posterior knob-like projection. The projec¬ tion itself could be thè contribution of thè exoccipital to thè formation of thè tripartite occipital condyle that is characteristic of squamates (with thè exception of uro- peltines). But thè generai shape and structure of thè bone
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON
fragment is such that no unequivocal and satisfactory interpretation can be offered.
Lateral to thè weak lateral extension of thè right ptery- goid sits a globular fragment of bone that has a counter- part on thè left side of thè skull and that could easily be mistaken for a massively developed lateral (transverse) process of thè pterygoid. However, these bone fragments are here interpreted as part of thè prootic. The description will focus on thè right element which is more fully exposed (Fig. 10). The fragment has an irregular appear- ance. As preserved, thè Teff side of thè fragment is of a bulbous structure, with a strongly convex surface and a moderate posterior projection that terminates in a blunt tip. This part of thè fragment formed a lappet that project- ed laterally from thè base of thè laterosphenoid-region of thè braincase and extended backwards to underlap thè posterior trigeminal (mandibular branch) foramen. Such a laterally projecting prootic (laterosphenoid) lappet is known from several basai macrostomatans such as boas and pythons (Rieppel et al., 2002), and it is also promi- nently developed in Haasiophis (Rieppel et al., 2003). Its presence, in fact, indicates thè presence of a laterosphe¬ noid, which is an alethinophidian feature (e.g., Rieppel, 1988). The part of thè fragment which lies topologically mediai (morphologically dorsal) to thè bulbous projection exposes a distinctly concave surface that is deeper anteri- orly than posteriorly. The anterior margin is also evenly curved (concave). This part of thè fragment formed thè posterior margin of thè mediai entry foramen into thè trigemino-facialis chamber (Rieppel, 1979). The concave surface behind thè entry foramen into thè trigemino- facialis chamber is thè part of thè prootic lying deep to thè exiting mandibular branch of thè trigeminal nerve: this is why it is more deeply concave anteriorly than posteriorly. There is one caveat with this interpretation, however. In basai macrostomatans such as Boa or Python , thè hyomandibular branch of thè facialis nerve typically pierces thè prootic within thè recess for thè exit of thè mandibular branch of thè trigeminal nerve, but a corre - sponding foramen cannot be located in thè same position in thè prootic fragment of specimen MSNM V 4014. However, thè facialis nerve pierces thè prootic in a more posterior position in anilioid snakes such as Anilius and Cylindrophis (Rieppel, 1977).
Both mandibular rami are preserved, but in spite of their superficially rather complete appearance, they offer only limited anatomical detail. The anterior tips of thè den- taries are smooth and rounded, indicating thè absence of a lower jaw symphysis. MeckeTs canal opens in front of thè splenial along thè medioventral edge of thè dentary (Fig. 10). The splenial is identifiable at thè ventral margin of thè mediai aspect of thè right mandibular ramus. A notch in its posterior part may indicate thè posterior mylohyoid fora¬ men (Fig. 10). If a bone fragment on thè mediai side of thè right mandibular ramus is correctly interpreted as thè ante¬ rior part of thè splenial, it suggests that thè splenial extended relatively far anteriorly. This is a macrostomatan feature that is also observed in Haasiophis (Rieppel et al., 2003). The exact configuration and posterior extent of thè posterior dentigerous process of thè dentary cannot be
established. The exact configuration of thè coronoid process is again impossible to establish. As described above, specimen MSNM V 3661 shares with Pachyrhachis a very prominent coronoid process of more or less rectangular outlines. Specimen MSNM V 4014 shows a weak elevation with a smoothly rounded dorsal margin in thè region behind thè posterior dentigerous process of thè right dentary. This elevation is recognized as a dorsal projection of thè right compound bone, which on its mediai side carries a facet for thè reception of thè coro¬ noid. The latter must have separated from thè mandibular ramus when thè slab and counterslab were split apart. An element on thè left side of thè skull, located at thè postero- lateral margin of thè preserved part of thè basisphenoid and narrowly overlapping thè latter, could well represent thè detached and dislocated left coronoid (Fig. 10). If cor¬ rectly identified, it would indicate that specimen MSNM V 4014 shares thè same tali coronoid as specimen MSNM V 3661. Both mandibular rami show thè compound bone in articulation with thè quadrates, and both display thè short, ‘knobby’ retroarticular process that is also observed in specimen MSNM V 3661.
The teeth are of similar structure in both specimens, a distinct ridging or fluting of thè enamel surface also being absent in MSNM V 4014.
A total of 162 vertebrae are preserved in specimen MSNM V 4014, with thè posterior tip of thè tail con- cealed by a coll of thè body. The anterior precloacal ver¬ tebrae again carry thè slender, bladelike neural spines that ascend in a posterodorsal direction from thè posterior part of thè neural arch (Figs. 3B, 7B). The first 30 vertebrae are preserved and exposed in lateral view, presumably again because of tilting caused by thè tali neural spines. Behind thè 30,h vertebra, thè neural spines are seen to decrease in height; a neural spine is present but relatively short in thè 37th preserved vertebra. Behind that element, thè vertebrae are split horizontally due to thè separation of thè part and counterpart slabs. Nevertheless, it can be stated that as in specimen MSNM V 3661, there are more than 37 anterior vertebrae that carry a distinct neural spine in specimen MSNM V 4014.
Fig. 1 1 - Precloacal vertebrae of specimen MSNM V 4014 as preserved in dorsal view.
18
OLIVIER RIEPPEL & JASON J. HEAD
Some of thè anterior precloacal vertebrae of specimen MSNM V 4014, preserved in lateral view, show a short and blunt yet distinct protuberance at thè posteroventral margin of thè centrimi (Fig. 7B), resembling thè rudimen- tary hypapophyses that were described for Simoliophis by Nopcsa (1925). Such rudimentary hypapophyses are absent on thè first 16 vertebrae. If present on thè 17th ver¬ tebra, it would be concealed by a body coil. The 18,h ver¬ tebra is poorly preserved but a small protuberance is present on thè posteroventral margin of thè centrum of thè 19,h through thè 23rd vertebrae. Behind thè 23rd vertebra, preservation becomes poor again, and behind thè 30lh ver¬ tebra, torsion of thè vertebral column begins. But thè rudimentary hypapophyses can be identified up to thè 30*, perhaps even 34,h, vertebra. The para-diapophysis (synapophysis) is situated very low on thè centrum towards its anterior end facing lateroventrally (Fig. 3B). The subcentrai foramen is located lateroventrally between thè synapophysis and thè rudimentary hypapophysis (where present). Finally, thè accessory prezygapophyseal process is absent in specimen MSNM V 4014.
Behind thè 37* vertebra, most of thè vertebrae are split horizontally, some above, and some below thè level of thè pre- and postzygapophyseal articulations. However, in thè posterior third of thè precloacal skeleton thè verte¬ brae are well preserved again and exposed in dorsal view. In thè posterior precloacal region, thè neural spine forms but a low ridge at thè anterior margin of thè neural arch. It increases slightly in height towards thè posterior mar¬ gin of thè neural arch, from where it projects in a pos- terodorsal direction, forming a short stubby process that overlaps thè anterior margin of thè neural arch of thè suc- ceeding vertebra. In cases where this short, posteriorly projecting neural spine is broken off (e.g., in thè 1 14* and 116* preserved vertebra) thè zygosphenal lamina (zygosphenal tectum: Rieppel et al., 2002) of thè suc- ceeding vertebra (thè 1 15* and 1 17* preserved element) is exposed in dorsal view. The zygosphenal tectum has a straight (117* vertebra) or only very slightly notched (115* vertebra, Fig. 1 1) anterior margin, unlike thè verte¬ bra described and illustrated by Rage & Escuillié (2000), which shows an anteriorly notched zygosphenal lamina. The 115* vertebra displays nicely thè zygosphene - zygantrum articulation, especially on thè left side. The zygosphene faces laterally, thè zygantrum medially, in a more or less vertically oriented piane of articulation. The zygantrum is formed by a pedicel that is set off from thè mediai side of thè prezygapophysis of thè preceding ver¬ tebra. The well-preserved vertebrae of thè posterior pre¬ cloacal region also show thè ‘swelling’ or protuberances on thè dorsal surface of thè neural arch on both sides of thè neural crest / spine (Fig. 11), which Rage & Escuillié (2000) found to be diagnostic of their new taxon, and which is thè only difference by which its vertebrae could be distinguished from those of Simoliophis. However, thè neural arch morphology as described for specimen MSNM V 4014 is also closely similar to that described for Pachyophis (Nopcsa, 1923a; Lee et al, 1999: 515, and fig. 1).
Nopcsa (1923a) described an interesting pattern of
variation of thè piane of articulation between thè pre- and postzygapophyses within thè precloacal vertebral column of Pachyophis. In thè anterior precloacal region, thè piane of articulation is weakly inclined (20° to 25° according to Lee et al., 1999; given its preservation, it seems doubtful that thè zygapophyseal angle can be estimated to an accu¬ rate degree in Pachyophis). In thè mid-precloacal region, thè articular surfaces on thè pre- and postzygapophyses are almost horizontally oriented. But in thè posterior pre¬ cloacal region, thè piane of articulation becomes progres- sively inclined again, thè prezygapophysis facing upwards and inwards, thè postzygapophysis facing down- wards and outwards. The swelling on thè dorsal surface of thè neural arch that is prominently expressed in thè mid- precloacal region recedes as thè vertebrae become rela- tively narrower in thè posterior precloacal region, until only deep grooves persist on either side of thè low and posteriorly projecting neural spine. This shows that thè swelling on thè dorsal surface of thè neural arch, which Rage & Escuillié (2000) took to be diagnostic for Eupodophis, is correlated with pachyostosis, as it is in specimen MSNM V 4014.
In specimen MSNM V 4014, thè first 16 vertebrae are well exposed in lateral view. A slight disarticulation between thè 7* and 8* vertebrae shows thè piane of articu¬ lation between thè prezygapophysis (facing upwards and inwards) and thè postzygapophysis (facing downwards and outwards) to be slightly inclined by about 20° to 25°, i.e., to a degree comparable to Pachyophis (Lee et al., 1999). The 1 9* through thè 24* vertebrae are tightly articulated, mak- ing it difficult to assess thè orientation of thè intervertebral articulation. The vertebrae following thè 24* element are crushed, and those behind thè 38* element are split horizon¬ tally, which renders it impossible to assess thè exact orien¬ tation of thè piane of articulation between thè pre- and postzygapophyses. However, within thè series of horizon¬ tally split vertebrae, thè 48* and 49* elements preserve their neural arch, and show thè piane of articulation between thè pre- and postzygapophyses to be only weakly inclined, per¬ haps by 15° to 20°. However, thè tight articulation between thè vertebrae renders that assessment somewhat tentative. Between thè 75* and 90* vertebrae, thè articulation between pre- and postzygapophysis is nearly horizontally oriented, inclined by no more than 5°. Behind thè 95* vertebra, thè inclination of thè articular surfaces of thè pre- and postzy¬ gapophyses rapidly increases again, as thè vertebrae become relatively narrower, and thè swelling on thè dorsal surface of thè neural arch recedes (due to a diminishing degree of pachyostosis) leaving deep grooves on either side of thè low neural spine. Around thè 125* vertebra, thè incli¬ nation of thè articular piane has increased to about 60°, thè prezygapophysis facing upwards and inwards, thè postzy¬ gapophysis facing downwards and outwards. Towards thè 145* vertebra, this inclination of thè articular piane becomes even more accentuateci, as does thè narrowing of thè vertebrae in generai. Specimen MSNM V 4014 thus documents a closely similar pattern of variation of thè inter¬ vertebral articulation between Eupodophis and Pachyophis. |
The relation of thè length to thè width of thè centrum, which is an indicator of thè degree of pachyostosis, is
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE &
hard to determine in specimen MSNM V 4014 given its preservation in dorsal view. The extent of pachyostosis along thè precloacal vertebral column is therefore diffi- cult to assess, but approximate measurements indicate that some degree of pachyostosis is expressed between thè 40,h and 155,h vertebrae, and that pachyostosis is most distinctly expressed between thè 65,h and 95th vertebrae.
Throughout thè precloacal region, thè para-diapoph- ysis (synapophysis) is set low on thè vertebral centrimi and hence is not exposed in dorsal view. It is exposed in a few vertebrae that have been split horizontally at a rela- tively low level, and these vertebrae, as well as thè posi- tion of thè proximal ends of thè ribs, indicate that thè synapophysis is again located towards thè anterior end of thè centrum. The proximal articular ends of thè ribs are generally buried under thè exposed dorsal part (neural arch) of thè vertebrae, in which case it is impossible to assess thè presence or absence of a tuberculum. Some posteriormost dorsal ribs are fully exposed, however, but no tuberculiform process can be identified on them. The ribs are curved in their proximal shoulder region but straight distally, indicating a lateral compression of thè body in Eupodophis.
The longest ribs in specimen MSNM V 4014 reach a length of 17 mm in thè middle precloacal region (75th ver¬ tebra; 16.5 mm for thè rib associated with thè 95th verte¬ bra). Specimen MSNM V 4014 resembles specimen MSNM V 3661 and Pachyrhachis, but not Pachyophis (or Haasiophis) in that elongated ribs extend further back- wards towards thè cloacal (i.e., hind limb) region, followed by two to four abruptly shortened ribs. The rib that articu-
ESCUILLIE. FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON ] 9
lates with thè 137* vertebra in specimen MSNM V 4014 equals thè length of 3.5 adjacent centra (approx. 5 mm). Abruptly much shortened ribs are associated with thè ver¬ tebrae 138 through 140. No rib is associated with thè 14T' vertebra. The left ilium, along with two tiny and rounded specks of bone, lies to thè left side of thè 1 4 1 st and 1 42nd vertebrae. Two stender strips of bone are emerging from below thè 145* and 146* vertebrae on their left side, but their identity cannot be unequivocally assessed. They do not appear to be ribs (no ribs are preserved on thè right side of thè vertebral column behind thè 140* vertebra), nor are they bifurcated as lymphapophyses would be.
The limb of specimen MSNM V 4014 is located in thè area of thè 141st to 144* (145*) preserved vertebrae (Fig. 8B). The vertebrae adjacent to thè limb and thè pelvic gir- dle rudiments show no trace of transverse processes or lymphapophyses. Indeed, none of thè specimens of Eupodophis show any indication of thè presence of trans¬ verse processes or lymphapophyses in thè cloacal region, and none of them allows thè identification of one or sev- eral putative ‘sacrai’ vertebra(e).
In specimen MSNM V 4014, thè orientation of thè articular surfaces of thè pre- and postzygapophyses becomes almost vertical in thè tail region (Fig. 12). Beginning with thè 148* vertebra, a distinct neural spine with rectangular outlines is developed, which rises from thè posterior part of thè neural arch and slightly slants in a posterodorsal direction. This results in an increasing height of thè short, laterally compressed tail.
The 145* through 148* vertebrae again develop a dis¬ tinct, ventrally projecting sagittal keel on thè ventral sur-
Fig. 12 - The cloacal and proximal tail region of thè vertebral column as preserved in specimen MSNM V 4014.
20
OLIVIER RIEPPEL & JASON J. HEAD
face of thè centrimi. At thè anterior end of thè centrimi of these four vertebrae, this keel is developed into a slender, blade-like and curved (anteriorly convex, posteriorly con¬ cave) process that projects in an anteroventral direction. More posteriorly (identifiable on thè 149lh through thè 154th vertebra), this anteroventral process is reduced to forni a short, medioventral (sagittal, unpaired) pedicel at thè anteroventral niargin of thè centrimi for thè articula- tion of thè chevrons. The chevrons ‘embrace’ this antero- medio-ventral pedicel with their short, proximally bifur- cating ‘arms’. The first vertebra to carry thè curved, anteroventrally projecting process is thè 146th preserved element in specimen MSNM V 3661, but thè 145lh verte¬ bra in specimen MSNM V 4014. This may very well be due to individuai variation. However, it also should be noted that thè atlas is not unequivocally identifiable behind thè skull of specimen MSNM V 4014, and a body coil conceals thè vertebrae following thè 16th preserved element. Conservatively, a single vertebra has been assumed to fili a gap in thè exposure of thè vertebral col- unin in this region. Considering these caveats, thè verte¬ bral counts given for specimen MSNM V 4014 through- out this description might be too low by one or perhaps even two elements.
The anteriormost chevron is associated with thè 149* vertebra (Fig. 12); it is somewhat shorter (though not much, as compared to specimen MSNM V 3661) than thè succeed- ing chevrons. The chevrons reach their maximum length at thè 153rd vertebra, and then decrease again in length towards thè tip of thè tail. The latter is concealed by a coil of thè body. The chevrons are tilted in a posteroventral direction relative to thè long axis of thè vertebral column.
A rod-shaped, somewhat curved element lies between thè fernur and thè vertebral column, its distai end slightly anterior but in dose proximity to thè proximal end of thè femur. This element is interpreted as thè ilium (Fig. 8B). Its length is 2.4 mm. Between thè ilium and thè proximal end of thè femur there are two flakes of bone which might represent remnants of other pelvic girdle elements not preserved in their entirety. Two similar specks of bone are also preserved near thè left ilium, which itself lies in dose proximity to thè vertebral column. There is no well- defined contact between thè ilium and any vertebrae in thè cloacal region.
The length of thè femur of specimen MSNM V 4014 equals thè length of 2.5 adjacent vertebral centra (Fig. 8B). The length of thè femur equals thè length of three adjacent vertebral centra in MSNM V 3661, and of 2.5 centra again in thè much larger specimen described by Rage & Escuillié (2000). The absolute length of thè femur is 3.46 mm in specimen MSNM V 4014 (average of three measurements). As in specimen MSNM V 3661 it is a straight, rather featureless bone with slightly expanded proximal and distai ends and a slightly constricted diaph- ysis. The tibia appears to be of a distinctly more robust structure than thè fibula, but a rib that lies between tibia and fibula may conceal some parts of thè latter (Fig. 8B). The length of thè tibia is 1 .8 mm, and it again is a straight element with slightly expanded ends and a slightly con¬ stricted diaphysis. The exact length of thè fibula cannot be measured, but thè element seems to be weakly curved, which would result in a spatium interosseum between tibia and fibula. No autopodal elements are preserved or present.
DISCUSSION
Specimens MSNM V 3661 and 4014 lend additional support to thè earlier finding that Eupodophis shares macrostomatan affinities (Rieppel et al., 2002). The poor- ly expressed articulation of thè right prefrontal with thè right frontal, as well as thè indication of thè presence of mediai frontal flanges on thè left frontal of specimen MSNM V 3661 (Fig. 6) suggest thè presence of a proki- netic joint. The supratemporal terminates in a slightly expanded, free ending posterior process. The vomer is slender and pointed posteriorly. The palatine carries a well-developed anterior dentigerous process, and thè slender mediai (choanal) process is located towards thè posterior end of thè bone. The pterygoid meets thè pala¬ tine in a characteristic tongue-and-groove articulation. The prootic flange underlapping thè posterior (mandibu- lar branch) trigeminal foramen indicates thè presence of a laterosphenoid. The basicranium is distinctly crested, sug- gesting thè presence of a well-developed constrictor inter- nus dorsalis musculature. The splenial reaches far for- wards on thè mediai aspect of thè lower jaw. All of thè above are characters that diagnose alethinophidian or macrostomatan snakes (Tchernov et al., 2000).
A character that has figured prominently in discus- sions of thè phylogenetic relationships of all Cenomanian snakes with well-developed hind limbs is thè identifica-
tion of a jugal in Pachyrhachis by Caldwell & Lee (1997; Lee & Caldwell, 1998; see also Lee, 1998). That identifi- cation was based on thè shape and relative shortness of thè element, and its position at thè ventral margin of thè orbit almost entirely in front of thè ventral tip of thè pos- torbital. Rieppel & Zaher (2000; see also Zaher & Rieppel, 2002) disputed this identification, and identified thè element in question as an incomplete ectopterygoid, broken across thè posterior end of thè maxilla in Pachyrhachis. However, specimen MSNM V 3661 of Eupodophis shows thè same element of same relative size and proportions and in an equivalent position, i.e., most- ly in front of thè ventral tip of thè postorbitai as is observed in Pachyrhachis. Careful preparation of thè left element in thè skull of specimen MSNM V 3661 (by C. Dal Sasso) revealed complete naturai edges around thè exposed parts of thè bone, thus refuting thè claim made by Rieppel & Zaher (2000) that thè equivalent element in Pachyrhachis was subject to breakage. However, Pachyrhachis shares with Eupodophis unusually elongat- ed frontals. If such elongated frontals are properly accounted for in a reconstruction of thè skull of Eupodophis (Fig. 13), it becomes apparent that thè elon¬ gated frontals carry thè postorbitai (articulating with thè anterolateral wings of thè parietal, and perhaps thè poste-
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON 2 1
riormost aspect of thè frontal) backwards. The elongated frontals are thus seen to be responsible for thè position of thè ectopterygoid mostly in front of thè ventral tip of thè postorbitai, which explains thè reason why these elements had initially been identified as jugals (Lee & Caldwell, 1998). Indeed, specimen MSNM V 3661 shows that if thè corresponding elements were identified as jugals, ectopterygoids would have to be absent.
The vertebral column of Eupodophis is very closely similar to that of Simoliophis, Pachyophìs and Mesophis (see systematic discussion below). The configuration of thè ribs indicates a lateral compression of thè body. Specimens MSNM V 3661 and 4014 show a very wide neural canal with a diameter that exceeds thè diameter of thè vertebral centrum. In contrast, thè specimen described by Rage & Escuillié (2000) is much larger than these specimens, and presents a ‘normal’ neural canal diameter where it is preserved. For that reason, we interpret thè wide neural canal in specimens MSNM V 3661 and 4014 as ontogenetic variation.
Eupodophis is unique among snakes in lacking lym- phapophyses in thè cloacal region. There are also no transverse processes present that would allow thè identi- fication of a ‘sacrai’ region. Small and rod-shaped pelvic girdle rudiments are present, but there is no evidence for a contact of thè ilium with thè axial skeleton. The most complete hind limb among all MSNM specimens of Eupodophis is preserved in MSNM V 4014 (Fig. 8B), where it consists of a femur, a tibia and a fibula (indica- tions of more distai elements are preserved in thè speci¬ men described by Rage & Escuillié, 2000). As is also thè case in Pachyrhachis and Haasiophis, thè femur in Eupodophis is a rather featureless, slender and straight element that lacks thè complex morphology which char-
acterizes thè femur of adult males of extant snakes that retain hind-limb rudiments (Kley et al., 2002). It is possi- ble that thè simple femur morphology in these marine snakes is a result of skeletal paedomorphosis, or that thè femur morphology of males of extant snakes is highly derived.
The character of Eupodophis that has attracted thè greatest attention is thè presence of elements that resem- ble chevrons in thè short tail (Figs. 8 A, 12). It is evident, however, that thè structure of thè caudal vertebrae and ‘chevrons’ of Eupodophis do not correspond to a non- ophidian or, in particular, varanoid morphology. In vara- noids, thè caudal centra bear paired ventral pedicels, located at thè posterior margin of thè vertebral centrum, with which thè chevrons articulate. These are Y-shaped elements with a short distai stem. In Eupodophis, thè chevrons (with thè possible exception of thè first two in thè series) have a comparatively much longer distai stem. Their proximal bifurcation results in thè formation of two short proximal ‘arms’ that embrace an unpaired, blade- like pedicel that is located at thè anterior end of thè cen¬ trum on thè midiine of its lower surface. The four verte¬ brae that precede thè first one associated with chevrons show a midventral crest which, at thè anterior end of thè centrum, extends into an anteroventrally curving, blade- like process. The formation of thè pedicels that articulate with thè chevrons result from thè reduction of this anteroventral process. This morphology is unique lor Eupodophis among all known squamates, and it creates problems of homology for thè interpretation of thè caudal vertebrae in other snakes.
The chevrons in non-ophidian squamates are consid- ered to represent intercentra, and they are located in an intercentral position in thè plesiomorphic condition. In
22
OLIVIER RIEPPEL & JASON J. HEAD
varanoids, thè chevrons become associated with thè pos- terior end of thè preceding vertebrae. These develop paired pedicels from thè posterior end of thè ventral sur- face of thè centra, which articulate with thè chevrons and which are considered genuine parts of thè centrum.
In snakes other than Eupodophis, thè anterior precloa- cal vertebrae develop unpaired ventral processes or hypa- pophyses from thè posterior part of thè centrum. On thè basis of some embryological evidence (Mànner, 1899: 65; Hoffstetter & Gasc, 1969: 284), these hypapophyses are considered to be homologous to intercentra that have become associated with thè posterior end of thè preceding vertebra. The extent to which hypapophyses are devel- oped posteriorly is variable among snakes (Hoffstetter & Gasc, 1969). In some taxa, a more or less distinctly devel- oped medioventral hemal crest or ridge may be present on thè middle and/or posterior precloacal vertebrae. In thè caudal region, snakes develop paired ventral processes (hemapophyses, unpaired in thè Uropeltinae), which fuse with thè centrum (Mànner, 1899: 66; Briinauer, 1910: 15; Hoffstetter & Gasc, 1969: 285), and which are again con¬ sidered homologous to intercentra (presumably to thè proximal ‘arms’ of thè Chevron).
In Eupodophis there is at best a very weakly devel- oped indication of a hypapophysis in thè anterior precloa¬ cal vertebrae. The ventral surface of thè trunk vertebrae is characteristically tripartite (Nopcsa, 1925) due to thè presence of subcentrai grooves that extend along a medi¬ ai prominence, a structural feature that is shared by Mesophis (Bolkay, 1925) and that is variably present in alethinophidians. In thè cloacal region, four vertebral centra develop a low medioventral keel from thè anterior end of which projects a curved and blade-like medioven¬ tral process. This keel and process could be considered an elaboration of thè hemal keel, or a peculiar transforma- tion of an unpaired hemapophysis. Again, hemapophyses
are described as undergoing ontogenetic fusion with thè centrum, which is why they are homologized with inter¬ centra associated with thè posterior end of thè preceding centrum. In thè fossil taxon Eupodophis, there is natural- ly no evidence for thè ontogenetic fusion of these process¬ es that project anteroventrally from thè anterior margin of thè centrum. Given their location at thè anterior end of thè centrum, it seems therefore more likely that these anteroventral processes present in four vertebrae of thè cloacal region represent genuine neomorphic formations of thè centrum. More posteriorly, within thè tail itself, these processes become reduced to small, blade-like and unpaired pedicels located on thè midiine of thè anterior margin of thè ventral surface of thè centra. With these articulate thè chevrons, which, if interpreted as inter¬ centra, would therefore have become associated with thè anterior end of thè posteriorly following centrum (rather than with thè posterior end of thè preceding centrum as in varanoids). Such an association of thè chevrons is unknown among non-ophidian squamates. Only some members of Agamidae show a slight posterior dislocation of thè chevrons “to a position under thè following verte¬ bra” (Hoffstetter & Gasc, 1969: 267). Otherwise, chevrons retain an intervertebral position or undergo an anterior dislocation amongst non-ophidian squamates.
Rieppel et al. (2002) showed that coding Eupodophis (and Wonambi ) for thè presence of chevrons did not alter their phylogenetic relationships with basai macrostom- atans. The presence of chevrons is, after all, a plesiomor- phic feature. And whereas Eupodophis is currently thè only snake known, fossil or extant, that shows thè pres¬ ence of chevrons, these certainly do not correspond to a plesiomorphic condition as indicated by out-group com- parison with non-ophidian squamates such as varanoids, and thè correlation of chevrons with thè apomorphic ante¬ rior pedicel in caudal vertebrae.
SYSTEMATIC CONCLUSIONS
In his originai description of Pachyrhachis problem- aticus, Haas (1979) noted thè similarity between thè ver¬ tebrae of this taxon and those of Simoliophis, and there¬ fore included Pachyrhachis in Simoliophidae (he attrib- uted thè family name to Nopcsa, 1924 [for 1925], who used Symoliophidae). Since he did not believe in thè ophidian status of Pachyrhachis, he removed thè simolio- phids from thè “snakes proper,” and referred thè family to snake-like “varanids” instead (Haas, 1979: 64). Similarly, Rage & Escuillié (2000) noted thè dose resemblance of thè precloacal vertebrae of Eupodophis with those of Simoliophis (see also Rage & Escuillié, 2003). Lee et al. (1999: 517) referred both Pachyophis and Pachyrhachis to Pachyophiidae (an emendation of Pachyophidae Nopcsa, 1923a; Pachyophinae in Nopcsa, 1923b; Pachyophiidae first appears in Bolkay, 1925: 130), noting that other fossil snakes such as Simoliophis may have to be referred to that family as well. Bolkay (1925) had already included Mesophis in Pachyophiidae.
Nopcsa (1923a: 139) first erected thè family Pachyo¬
phidae to include Pachyophis (Nopcsa, 1923b: 124, used Pachyophinae as a subfamily of Cholophidae). Following Cannatella’s (1990) orthographic analysis, Lee et al. (1999: 510) “more correctly” applied thè “taxon name Pachyo¬ phiidae to Pachyophis, Pachyrhachis, and all taxa more closely related to these genera than to modern snakes” (Lee et al, 1999: 517). In thè case of Pachyophis thè question is what thè proper stem of ‘ophis’ would be? The problem is that ‘ophis’ has a peculiar genitive in Attic Greek. According to some sources, ‘opheos’ would be thè correct genitive, while in other dialects thè genitive of ‘ophis’ is straightforwardly ‘ophios’, which gives thè stem ‘ophi-‘. According to thè Code of Nomenclature, this latter genitive translates into thè family name Ophiidae. On thè other hand, Ophidia is thè neuter plural of a genuine Greek diminutive ophidion (‘little snake’). This derived noun is built on thè stem oph- plus thè diminutive suffix -idion. This argues for Ophidae. So, it is arguable that Pachyo¬ phidae is not ‘incorrect Greek,’ but it is also arguable that Pachyophiidae follows thè rules of thè Code more straight-
NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIE, FROM THE CENOMANLAN (LATE CRETACEOUS) OF LEBANON
forwardly (Howard Don Cameron, personal communica- tion; see also Cannatella, 1990).
In his description of thè Simoliophis material from Egypt, Nopcsa (1925) recognized a dose afifinity of Pachyophis and Simoliophis, raising thè question whether thè two taxa should be treated as representatives of two families (Pachyophidae and Symoliophidae [s/c] respec- tively), or as two separate subfamilies within one family. He later (Nopcsa, 1928) formally accepted thè latter solu¬ tion, listing Pachyophinae and Symoliophinae as subfam¬ ilies of his Cholophidae. Hoffstetter (1955) lists Pachyophidae and Simoliophidae as two separate families within Cholophidia. Romer (1956: 563) includes Pachyophis and Simoliophis in thè subfamily liSimolio- phinae ( Pachyophinae )” that is included in his family Palaeophidae (which he equates with Cholophidae and Cholophidia). Kuhn (1967) finally considered as valid names Simoliophidae Hoffstetter (1955: 464), and Pachyophinae Nopcsa (1923b: 124).
In his discussion of thè material attributed to Simoliopohis, Nopcsa (1925) found thè vertebrae to be variable to a degree that only one species name could be justified, which by priority is Simoliophis rochebrunei Sauvage, 1880 (see also Rage, 1984). Since that time, another species, Simoliophis libycus was described by | Nessov et al. (1998), but they misidentified thè anterior precloacal vertebrae for posterior precloacal elements, which removes their main reason for thè recognition of a > second species. Rage & Escuillié (2003: 7) consider thè validity of Simoliophis libycus questionable on thè grounds that thè characters that differentiate this taxon from Simoliophis rochebrunei might be due to ontogenet- ic variation or variation along thè vertebral column. Rage & Escuillié (2003: 7) furthermore consider thè material described by Nopcsa (1925) to comprise two different taxa, one of which is a separate species of Simoliophis that remains to be named. Caldwell & Lee (1997; see also Lee & Caldwell, 1998) correctly recognized Ophiomor- phus colberti Haas, 1980b [. Estesius , Wallach, 1984], as a I: subjective junior synonym of Pachyrhachis problemati- 1 cus, and Head (J. Head, in preparation) recognizes thè synonymy of Pachyrhachis with Simoliophis, a conclu- sion that was also suggested by Rage & Escuillié (2003: 7). Closely similar forms are Pachyophis (Nopcsa, 1 923a) and Mesophis (Bolkay, 1925). However, given thè many characters of skull structure, as well as differences in postcranial anatomy (lack of tali neural spines in thè ante¬ rior precloacal region, configuration of thè rib cage, dif- ferentiation of mid-precloacal pachyostosis), there can be no doubt that Haasiophis (Tchemov et al., 2000; Rieppel et al., 2003) represents a separate genus, yet related to thè above mentioned taxa and again included in Pachyo- phiidae (see also Rieppel et al., 2002). The same case is . not quite as easily made for Eupodophis.
Rage & Escuillié (2000) recognized thè dose similar- ity of thè vertebrae of Eupodophis to those of Simoliophis, and cited thè protuberances on thè dorsal surface of thè neural arch as thè only difference from thè latter genus (these protuberances are correlated with pachyostosis as described above; see also Fig. 1 1). At thè
same time, Rage & Escuillié (2003: 7) find thè dorsal tubercles on thè vertebrae of Eupodophis to preclude thè latter genus from being a subjective junior synonym of Simoliophis, a synonymy which they consider possible for Pachyrhachis and Haasiophis (but see above). Specimen MSNM V 3660 is preserved in ventral view, and both in size and structure matches thè specimen described by Rage & Escuillié (2000) as well as thè material referred to Simoliophis (Nopcsa, 1925; Nessov et al., 1998). Specimens MSNM V 3661 and 4014 are much smaller than thè known material referred to Pachyrhachis and Simoliophis, and thè material referred to Eupodophis just mentioned, but they share characters both with Pachyrhachis (in thè skull) and Eupodophis (in thè post¬ cranial skeleton), and therefore they also share similari- ties, impliciter, with Simoliophis.
We are here taking a conservative approach to speci¬ mens MSNM V 3661 and 4014 in considering them juve- niles of Eupodophis descouensi Rage & Escuillié (2000), rather than a separate taxon of small snakes (see thè dis¬ cussion of thè unusually wide neural canal in these speci¬ mens above). One of thè problems here is that thè skull of thè holotype of Eupodophis descouensi is very poorly and incompletely preserved, and no skull is preserved in spec¬ imen MSNM V 3660. This leaves for comparison thè post¬ cranial skeleton only. It is not inconceivable, however, that future finds of larger specimens with a skull referable to Eupodophis descouensi may show that thè small individu¬ ate here described belong to a different species.
The skull characters which thè specimens MSNM V 3661 and 4014 collectively share with Pachyrhachis (but not with Haasiophis) include thè relatively long frontals, thè shape of thè postorbitai, thè shape and proportions of thè ectopterygoid, and thè high coronoid of rectangular outlines (thè quadrate is broad in MSNM V 4014, but nar- row in MSNM V 3661; this character must therefore be considered polymorphic unless it is due to a preservation- al artifact). The postcranial characters which thè speci¬ mens MSNM V 3661 and 4014 share with Eupodophis are thè absence of lymphapophyses in thè cloacal region (present in Pachyrhachis : see thè re-interpretation of thè putative sacrai rib described by Lee & Caldwell [1998] as a lymphapophysis [Lee, 1998; Scanlon & Lee, 2000]), and thè presence of elongated, proximally bifurcated chevrons (possibly absent in Pachyrhachis, but only thè very proxi- mal part of thè tail is known for that genus). The speci¬ mens MSNM V 3661 and 4014 ateo share characters with Haasiophis, such as thè posteriorly weakly expanded free ending process of thè supratemporal. Given this character distribution, there can be little doubt that specimens MSNM V 3661 and 4014 may represent juvenile speci¬ mens of Eupodophis, that Eupodophis may be more close¬ ly related to Pachyrhachis than Haasiophis on thè basis of skull structure, and that thè three taxa represent three dif¬ ferent species. However, no part of thè skull is preserved in thè material that has so far been referred to Simoliophis, nor is it known whether Simoliophis did have lym¬ phapophyses in thè cloacal region, or chevrons in thè tail. Since Pachyrhachis problematicus cannot be recognized as different from Simoliophis, Pachyrhachis can be con-
24
OLIVIER RIEPPEL & JASON J. HEAD
sidered a junior synonym of Simoliophis. In contrast, Eupodophis can be recognized as distinct, such that it is kept separate from Simoliophis. Haasiophis would then be a third, monotypic genus in thè family Pachyophiidae Nopcsa 1923 (1923a).
Amongst thè marine Cenomanian snakes here dis¬ cusseci, mid-trunk pachyostosis is expressed to a lesser degree in Eupodophis and Mesophis than in Haasiophis , Pachyrhachis , Pachyophis, Simoliophis. However, there is a possibility of ontogenetic differences in thè extent of pachyostosis. Pachyrhachis shares with Eupodophis thè abrupt shortening of thè last few dorsal ribs (not pre- served in Mesophis and Simoliophis ), thè elongated frontals, thè broad quadrate (conservatively considered polymorphic in Eupodophis) and thè tali and rectangular coronoid process (thè skull is not preserved in Mesophis
and Simoliophis). Pachyophis shares with Pachyrhachis, Eupodophis and Simoliophis thè tali neural spines on thè anterior precloacal vertebrae, and thè laterally strongly compressed body, thè latter a character that is also pres- ent in Mesophis. In summary, thè genera Eupodophis, Mesophis, Pachyophis, Pachyrhachis and Simoliophis appear more closely related to each other and to Simoliophis than either is to Haasiophis. This, indeed, is also strongly reflected in terms of skull morphology (as described and discussed above), which is known for Eupodophis, Haasiophis, and Pachyrhachis, but not for thè other three taxa of pachyostotic Cenomanian marine snakes. At thè bottom line, however, some of thè taxa (Mesophis, Pachyophis, and Simoliophis) are just too incompletely preserved to warrant a more complete phy- logenetic analysis.
GENERAL CONCLUSIONS
The genus Eupodophis from thè Cenomanian of Lebanon is part of a mid-Cretaceous radiation of marine snakes in thè western Tethyan (Neotethys) faunal realm, which also includes thè genera Haasiophis, Mesophis, Pachyophis, Pachyrhachis, and Simoliophis (Rage & Escuillié, 2003). Amongst those taxa, well-developed hind limbs are known to occur in Eupodophis, Haasiophis, and Pachyrhachis, but recognizing their dose interrelationships, Rage & Escuillié (2003) hypoth- esized that such hind limbs may also have been present in thè other genera known from that radiation. As noted by Rage & Escuillié (2003: 7; see also Rage & Escuillié, 2000), thè combination in these snakes of a macrostom- atan skull with thè presence of well-developed hind limbs “poses a serious problem,” because “two diametrically opposite” phylogenetic conclusions may obtain. The hypothesis that thè fossil snakes with well-developed hind limbs are basai to all extant snake lineages (Lee & Caldwell, 1998; Lee & Scanlon, 2002) implies that thè macrostomatan skull is primitive for snakes (Rage & Escuillié, 2000, 2003). Alternatively, and on thè basis of thè current morphology-based phylogeny for extant snake lineages (which places scolecophidians and ‘anilioids’ basai to macrostomatans: e.g., Cundall et al., 1993; Cundall & Greene, 2000; Tchernov et al., 2000; Lee & Scanlon, 2002), thè hypothesis that thè fossil snakes are basai macrostomatans (Rieppel et al., 2002) implies that thè well-developed hind limbs either redeveloped in thè fossil snakes, or that hind limbs were subject to repeated reduction and loss within snakes (Rieppel et al., 2003; see also thè discussion in Coates & Ruta, 2000).
However, recent molecular studies of snake interrela¬ tionships (Wilcox et al., 2002; Slowinski and Lawson, 2002; Vidal & David, 2004; Vidal & Hedges, 2004) have called into question thè current understanding of thè phy¬ logeny of extant snakes as based on morphology in several respects. Most importantly, there is a recurrent signal for a basai position of thè macrostomate genera Tropidophis and Tachyboa outside thè conventional macrostomatan clade (Wilcox et al., 2002), possibly dose to Anilius (Vidal & David, 2004; Vidal & Hedges, 2004). Conversely, thè non- macrostomate genus Cylindrophis and thè related uro- peltines may nest inside thè conventional macrostomatan clade (Wilcox et al., 2002; Vidal & David, 2004; Vidal & Hedges, 2004). These molecular findings, if confirmed by future studies, may well result in a novel understanding of snake interrelationships that may cast new light on thè interpretation of thè Cenomanian snakes with well-devel¬ oped hind limbs.
Acknowledgements
We thank Giorgio Teruzzi and Cristiano Dal Sasso for permission to work on thè specimens of Eupodophis in their care, and Cristiano Dal Sasso for important help and support in getting this project off thè ground. David Cundall, Maureen Kearney, and Nate Kley reviewed an earlier draft of thè paper, offering much appreciated advice and criticism. This research was supported by thè US National Science Foundation Grant DEB-0235628.
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Vidal N. & Hedges S.B., 2004 - Molecular evidence for a terrestrial origin of snakes. Proceedings of thè Royal Society of London, Biological Sciences (Suppl.), London, 271: S226-S229.
Vidal N. & David P., 2004 - New insights into thè early history of snakes inferred from two nuclear genes. Molecular Phylogenetics and Evolution, San Diego, 31:783-787/
Wilcox T.P., Zwickl D.J., Heath A. & Hillis D.M., 2002 - Phylogenetic relationships of thè dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, San Diego, 25: 361-371.
Zaher H., 1 998 - The phylogenetic position of Pachyrhachis within snakes (Squamata, Lepidosauria). Journal of Vertebrate Paleontology, Lawrence KA, 18: 1-3.
Zaher H. & Rieppel O., 2002 - On thè phylogenetic relation¬ ships of thè Cretaceous snakes with legs, with special reference to Pachyrhachis problematicus (Squamata, Serpentes). Journal of Vertebrate Paleontology, Lawrence KA, 22: 104-109.
Olivier Rieppel, Department of Geology, The Field Museum, 1400 S Lake Shore Drive, Chicago, IL 60605-2496, USA;
e-mail: rieppel@fieldmuseum.org
Jason J. Head, Department of Paleobiology, The National Museum of Naturai History, Smithsonian Institution,
PO. Box 37012, NHB MRC121, Washington, D.C. 20013-7012, USA; e-mail: head.jason@nmnh.si.edu
New specimens of thè fossil snake genus Eupodophis Rage & Escuillé, from Cenomanian (Late Cretaceous) of Lebanon Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXII - Fascicolo II
Volume XVII
- PINNA G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Alpe Turati (Erba, Como). Famiglie Lytoceratidae, Nannolytoceratidae, Hammatoceratidae (excl. Phymatoceratinae) Hildoceratidae (excl. Hildoceratinae e Bouleiceratinae). pp. 1-70, 2 tavv. n.t., 6 fìgg-, 6 tavv.
- VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi delI’Anisico superiore di Lenna in Val Brembana (Bergamo), pp. 71- 142, 5 fìgg-, 11 taw.
- PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Alpe Turati (Erba, Como). Generi Hildoceras, Phymatoceras, Paroniceras e Frechiella. Conclusioni generali, pp. 143-204, 2 figg.,
6 tavv.
Volume XVIII
- PINNA G., 1969 - Revisione delle ammoniti figurate da Giuseppe Meneghini nelle Tavv. 1 -22 della « Monographie des fossiles du cal¬ cane rouge ammonitique» ( 1 867- 1881). pp. 5-22, 2 Jìgg., 6 taw.
- MONTANARI L., 1969 - Aspetti geologici del Lias di Gozzano (Lago d’Orta). pp. 23-92, 42 fìgg.. 4 taw. n.t.
- PETRUCCI F„ BORTOLAMI G. C. & DAL P1AZ G. V., 1970 - Ricerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul suo substrato cristallino, pp. 93-169, con carta a colori al 1:40.000, 14 fìgg., 4 taw. a colori e 2 b.n.
Volume XIX
- CANTALUPPI G., 1970 - Le Hildoceratidae del Lias medio delle regioni mediterranee - Loro successione e modificazioni nel tempo. Riflessi biostratigrafici e sistematici, pp. 5-46, con 2 tabelle nel testo.
- PINNA G. & LEVI-SETTI F„ 1971 - I Dactylioceratidae della Provincia Mediterranea (Cephalopoda Ammonoidea) . pp. 47-136, 21 fìgg., 12 taw.
- PELOSIO G., 1973 - Le ammoniti del Trias medio di Asklepieion (Argolide, Grecia) - I. Fauna del «calcare a Ptychites » (Anisico sup. ).pp. 137-168, 3 fìgg., 9 taw.
Volume XX
- CORNAGGIA CASTIGLIONI O., 1971 - La cultura di Remedello. Problematica ed ergologia di una facies dell’Eneolitico Padano, pp. 5-80, 2 fìgg., 20 taw.
- PETRUCCI F. 1972 -Il bacino del Torrente Cinghio (Prov. Parma). Studio sulla stabilità dei versanti e conservazione del suolo, pp. 81- 127, 37 fìgg., 6 carte tematiche.
- CERETTI E & POLUZZI A., 1973 - Briozoi della biocalcarenite del Fosso di S. Spirito (Chieti, Abruzzi)./?/?. 129-169, 18 fìgg., 2 taw.
Volume XXI
- PINNA G., 1974 - I crostacei della fauna triassica di Cene in Val Seriana (Bergamo)./?/?. 5-34, 16 fìgg., 16 taw.
- POLUZZI A., 1975 - I Briozoi Cheilostomi del Pliocene della Val d’Arda (Piacenza, Italia)./?/?. 35-78, 6 fìgg., 5 taw.
- BRAMBILLA G., 1976 - I Molluschi pliocenici di Villalvemia Alessandria). I. Lamellibranchi. pp. 79-128, 4 fìgg., 10 taw.
Volume XXII
- CORNAGGIA CASTIGLIONI O. & CALEGARI G., 1978 - Corpus delle pintaderas preistoriche italiane. Problematica, schede, icono¬ grafia./?/?. 5-30, 6 fìgg., 13 taw.
- PINNA G., 1979 - Osteologia dello scheletro di Kritosaurus notabi¬ li (Lambe, 1914) del Museo Civico di Storia Naturale di Milano (Ornithischia Hadrosauridae). pp. 31-56, 3 fìgg.. 9 taw.
- BIANCOTTI A., 1981 - Geomorfologia dell’Alta Langa (Piemonte meridionale), pp. 57-104, 28 fìgg., 12 tabb., 1 carta f. t.
Volume XXIII
- GIACOBINI G„ CALEGARI G. & PINNA G., 1982 - 1 resti umani fossili della zona di Arena Po (Pavia). Descrizione e problematica di una serie di reperti di probabile età paleolitica./?/?. 5-44, 4 fìgg.. 16 taw.
- POLUZZI A., 1982 - I Radiolari quaternari di un ambiente idroter¬ male del Mar Tirreno, pp. 45-72, 3 fìgg., 1 tab., 13 taw.
- ROSSI F., 1984 - Ammoniti del Kimmeridgiano superiore Berriasiano inferiore del Passo del Furio (Appennino Umbro- Marchigiano)./?/?. 73-138, 9 fìgg., 2 tabb., 8 taw.
Volume XXIV
- PINNA G., 1984 - Osteologia di Drepanosaurus unguicaudatus, lepi- dosauro triassico del sottordine Lacertilia. pp. 7-28, 12 fìgg., 2 taw.
- NOSOTTI S., PINNA G., 1989 - Storia delle ricerche e degli studi sui rettili Placodonti. Parte prima 1830-1902./?/?. 29-86, 24 fìgg., 12 taw.
Volume XXV
- CALEGARI G., 1989 - Le incisioni rupestri di Taouardei (Gao, Mali). Problematica generale e repertorio iconografico./?/?. 1-14, 9 fìgg., 24 taw.
- PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia funziona¬ le e paleoecologia del rettile placodonte Psephoderma alpinum Meyer, 1858 . pp. 15-50. 20 fìgg., 9 taw.
- CALDARA R., 1990 - Revisione Tassonomica delle specie palearti¬ che del genere Tychius Germar (Coleoptera Curculionidae). pp. 51- 218, 575 fìgg.
Volume XXVI
I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia,
Placodontia). pp. 1-21, 23 fìgg.
II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara prei¬
storico: dati e interpretazioni./?/?. 25-556. 647 fìgg.
Ili - ANDRI E. e ROSSI F., 1993 - Genesi ed evoluzione di frangenti, cinture, barriere ed atolli. Dalle stromatoliti alle comunità di scoglie¬ ra moderne, pp. 559-610, 49 fìgg., 1 tav.
Volume XXVII
I - PINNA G. & GHISELIN M. edited by, 1996 - Biology as History. N.
1 . Systematic Biology as an Historical Science, pp. 1-133, 68 fìgg.
II - LEONARDI C. & SASSI D. a cura di, 1997 - Studi geobotanici ed
entomofaunistici nel Parco Regionale del Monte Barro./?/?. 135-266.
Volume XXVIII
I - BANFI E. & GALASSO G., 1998 - La flora spontanea della città di Milano alle soglie del terzo millennio e i suoi cambiamenti a partire dal 1700./?/?. 267-388.
Volume XXIX
I - CALEGARI G., 1999 - L’arte rupestre dell’Eritrea. Repertorio ragionato ed esegesi iconografica./?/?. 1-174.
Volume XXX
I - PEZZOTTA F. edited by, 2000 - Mineralogy and petrology of shal-
low depth pegmatites. Paper from thè First International Workshop. pp. 1-117.
II - PARISI B„ FRANCHINO A. & BERTI A. con la collaborazione di
POTENZA B. & RUBINI D., 2000 - La Società Italiana di Scienze Naturali 1855 - 2000. Percorsi storici./?/?. 1-163.
Ili - DE ANGELI A. & GARASSINO A., 2002 - Galatheid, chirostylid and porcellanid decapods (Crustacea, Decapoda, Anomura) from thè Eocene and Oligocene of Vicenza (N Italy). pp. 1-40.
Volume XXXI
I - NOSOTTI S. & RIEPPEL O., 2002 - The braincase of Placodus
Agassiz, 1833 (Reptilia, Placodontia)./?/?. 1-18.
II - MARTORELLI G., 2002 - Monografia illustrata degli uccelli di
rapina in Italia. ( 1 895). Riedizione a cura di Fausto Barbagli, pp. [XX] 1- 216.
Ili - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphargis dalsassoi n.gen.n.sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano (Lombardy, N Italy). pp. 1-33.
Volume XXXII
I - ALESSANDRELLO A., BRACCHI G. & RIOU B„ 2004 - Polychaete, sipunculan and enteropneust worms from thè Lower Callovian (Middle Jurassic) of La Voulte-sur-Rhóne (Ardèche, Frane e)./?/?. 1-16, 1 tav.
Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali Museo Civico di Storia naturale, Corso Venezia 55 - 20121 Milano Pubblicazione disponibile al cambio
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TOOÓL
MEMORIE
LIBRARY
JUN 7 2005
Volume XXXII - Fascicolo III HARVAR K. _ UNIVERSITY
GIACOMO BRACCHI & ANNA ALESSANDRELLO
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA)
AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS LAGERSTÀTTEN OF HAQEL, HADJULA AND AL-NAMOURA (LEBANON)
della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
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MILANO FEBBRAIO 2005
Elenco delle Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume I
I - CORNALIA E:, 1865 - Descrizione di una nuova specie dei genere
Felis: Felis jacobìta (Com.). 9 pp., 1 tav.
II - MAGNI-GRIFFI F., 1865 - Di una specie d 'Hippolais nuova per l’Ita¬
lia. 6 pp., 1 tav.
Ili - GASTALDI B., 1865 - Sulla riescavazione dei bacini lacustri per opera degli antichi ghiacciai. 30 pp., 2 figg., 2 tavv.
IV - SEGUENZA G., 1865 - Paleontologia malacologica dei terreni terziarii
del distretto di Messina. 88 pp., 8 tavv.
V - GIBELLI G., 1865 - Sugli organi riproduttori del genere Verrucaria,
16 pp., 1 tav.
VI - BEGGIATO F. S., 1865 - Antracoterio di Zovencedo e di Monteviale
nel Vicentino, 10 pp., 1 tav.
VII - COCCHI I., 1865 - Di alcuni resti umani e degli oggetti di umana industria dei tempi preistorici raccolti in Toscana. 32 pp., 4 tavv.
Vili - TARGIONI-TOZZETTI A., 1866 - Come sia fatto l'organo che fa lume nella lucciola volante dell’Italia centrale (Luciola italica) e come le fibre muscolari in questo ed altri Insetti ed Artropodi. 28 pp., 2 tavv.
IX - MAGGI L., 1865 - Intorno al genere Aeolosoma. 18 pp., 2 tavv.
X - CORNALLA E., 1865 - Sopra i caratteri microscopici offerti dalle Canta¬
ridi e da altri Coleotteri facili a confondersi con esse. 40 pp., 4 tavv.
Volume II
I - ISSEL A., 1866 - Dei Molluschi raccolti nella provincia di Pisa, 38 pp.
II - GENTILLI A., 1866 - Quelques considérations sur l’origine des bassins
lacustres, àpropos des sondages du Lac de Come. 12 pp., 8 tavv.
III - MOLON F., 1867 - Sulla flora terziaria delle Prealpi venete. 140 pp.
IV - D’ACHIARDI A., 1866 - Corallarj fossili del terreno nummulitico
delle Alpi venete. 54 pp., 5 tavv.
V - COCCHI I., 1866 - Sulla geologia dell’alta Valle di Magra. 18 pp., 1 tav.
VI - SEGUENZA G., 1866 - Sulle importanti relazioni paleontologiche di
talune rocce cretacee della Calabria con alcuni terreni di Sicilia e del¬ l’Africa settentrionale. 18 pp., 1 tav.
VH - COCCHI I., 1 866 - L’uomo fossile nell’Italia centrale. 82 pp., 21 figg., 4 tavv. Vili - GARO VAGLIO S., 1866 - Manzonia cantiana, novum Lichenum Angiocarporum genus propositum atque descriptum. 8 pp., 1 tav.
IX - SEGUENZA G., 1 867 - Paleontologia malacologica dei terreni terziarii
del distretto di Messina (Pteropodi ed Eteropodi). 22 pp., 1 tav.
X - DURER B., 1867 - Osservazioni meteorologiche fatte alla Villa Carlot¬
ta sul lago di Como, ecc. 48 pp. 11 tavv
Volume III
I - EMERY C., 1873 - Studii anatomici sulla Vipera Redii. 16 pp., 1 tav.
II - GARO VAGLIO S., 1867 - Thelopsis, Belonia, Weitenwebera et Limbo-
ria, quatuor Lichenum Angiocarporum genera recognita iconibusque illustrata. 12 pp., 2 tavv.
III - TARGIONI-TOZZETTI A., 1 867 - Studii sulle Cocciniglie. 88 pp., 7 tavv.
IV - CLAPARÈDE E. R. e PANCERI P., 1867 - Nota sopra un Alciopide
parassito della Cydippe densa Forsk. 8 pp. 1 tavv.
V - GARO VAGLIO S., 1871 - De Pertusariis Europae mediae commenta¬
no. 40 pp., 4 tavv.
Volume IV
I - D’ACHIARDI A., 1868 - Corallarj fossili del terreno nummulitico del- l’Alpi venete. Parte 1 1 . 32 pp. 8 taw.
II - GARO VAGLIO S., 1 868 - Octona Lichenum genera vel adhuc controver¬
sa, vel sedis prorsus incertae in systemate, novis descriptionibus iconibu¬ sque accuratissimis illustrata. 18 pp., 2 taw.
III - MARINONI C., 1868 - Le abitazioni lacustri e gli avanzi di umana
industria in Lombardia. 66 pp., 5 figg., 7 tavv.
IV - (Non pubblicato).
V - MARINONI C., 1871 - Nuovi avanzi preistorici in Lombardia. 28 pp.,
3 figg., 2 tavv.
NUOVA SERIE
Volume V
I - MARTORELLI G., 1895 - Monografia illustrata degli uccelli di rapina in Italia. 216 pp., 46 figg., 4 tavv.
Volume VI
I - DE ALESSANDRI G„ 1897 - La pietra da cantoni di Rosignano e di
Vignale. Studi stratigrafici e paleontologici. 104 pp., 2 tavv., I carta.
II - MARTORELLI G., 1898 - Le forme e le simmetrie delle macchie
nel piumaggio. Memoria ornitologica. 112 pp., 63 figg., 1 tavv.
Ili - PAVESI P., 1901- L’abbate Spallanzani a Pavia. 68 pp., 14 figg., 1 tav.
Volume VII
I - DE ALESSANDRI G., 1910 - Studi sui pesci triasici della Lombardia. 164 pp., 9 tavv.
Volume Vili
I - REPOSSI E., 1915 - La bassa Valle della Mera. Studi petrografici e
geologici. Parte l. pp. 1-46, 5 figg., 3 taw.
II - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi petrogra¬
fici e geologici. Parte IL pp. 47-186, 5 figg. 9 taw.
III - AIRAGHI C., 1917 - Sui molari d’elefante delle alluvioni lombarde,
con osservazioni sulla filogenia e scomparsa di alcuni Proboscidati. pp. 187-242, 4 figg., 3 taw.
Volume IX
I - BEZZI M., 1918 - Studi sulla ditterofauna nivale delle Alpi italiane, pp.
1-164, 7 figg. 2 tavv.
II - SERA G. L., 1 920 - Sui rapporti della conformazione della base del cra¬
nio colle forme craniensi e colle strutture della faccia nelle razze uma¬ ne. (Saggio di una nuova dottrina craniologica con particolare riguardo dei principali cranii fossili), pp. 165-262, 7 figg., 2 taw.
Ili - DE BEAUX O. e FESTA E., 1927 - La ricomparsa del Cinghiale nel¬ l’Italia settentrionale-occidentale,/?/?. 263-320, 13 figg., 7 taw.
Volume X
I - DESIO A., 1929 - Studi geologici sulla regione dell’Albenza (Prealpi
Bergamasche), pp. 1-156, 27 figg., 1 tav., 1 carta.
II - SCORTECCI G., 1937 - Gli organi di senso della pelle degli Agamidi.
pp. 157-208, 39 figg. 2 tavv.
Ili - SCORTECCI G., 1941- 1 recettori degli Agamidi. pp. 209-326, 80 figg.
Volume XI
I - GUIGLI A D., 1 944 - Gli Sfecidi italiani del Museo di Milano (Hymen.)
pp. 1-44, 4 figg., 5 taw.
II-III - G1ACOMIN1 V. e PIGNATTI S., 1955 - Flora e Vegetazione dell’Al¬ ta Valle del Braulio. Con speciale riferimento ai pascoli di altitudine.
pp. 45-238, 31 figg., 1 carta.
Volume XII
I - VIALLI V., 1956 - Sul rinoceronte e l’elefante dei livelli superiori della serie lacustre di Leffe (Bergamo), pp. 1-70, 4 figg. 6 taw.
I - VENZO S., 1957 - Rilevamento geologico dell’anfiteatro morenico del Garda. Parte I: Tratto occidentale Gardone-Desenzano. pp. 71-140, 14 figg., 6 tavv., 1 carta.
Ili - VIALLI V., 1959 - Ammoniti sinemuriane del Monte Albenza (Berga¬ mo), pp. 141-188, 2 figg., 5 taw.
Volume XIII
I - VENZO S., 1961- Rilevamento geologico dell’anfiteatro morenico del
Garda. Parte II. Tratto orientale Garda-Adige e anfiteatro atesino di Rivoli veronese, pp. 1-64, 25 figg., 9 tavv., 1 carta.
II - PINNA G., 1963 - Ammoniti dei Lias superiore (Toarciano) dell’Alpe
Turati (Erba, Como). Generi Mercaticeras, Pseudomercaticeras e Bro- dieia. pp. 65-98, 2 figg., 4 tavv.
III - ZANZUCCHI G., 1 963 - Le Ammoniti del Lias superiore (Toarciano) di En-
tratico in Val Cavallina (Bergamasco orientale), pp. 99-146, 2 figg., 8 taw.
Volume XIV
I - VENZO S., 1965 - Rilevamento geologico dell’anfiteatro morenico fron¬
tale del Garda dal Chiese all’Adige, pp. 1-82, 11 figg-, 4 taw., 1 carta.
II - PINNA G., 1966 - Ammoniti del Lias superiore (Toarciano) dell’Alpe
Turati (Erba, Como). Famiglia Dactylioceratidae. pp. 83-136, 4 taw.
III - DIENI I., MASSARI F. e MONTANARI L„ 1966 - Il Paleogene dei
dintorni di Orosei (Sardegna), pp. 13-184, 5 figg., 8 tavv.
Volume XV
I - CARETTO P. G., 1966 - Nuova classificazione di alcuni Briozoi plioce¬
nici, precedentemente determinati quali Idrozoi del genere Hydractinia Van Beneden. pp. 1-88, 27 figg. 9 tavv.
II - DIENI I. e MASSARI F., 1966 - Il Neogene e il Quaternario dei dintorni
di Orosei (Sardegna), pp. 89-142, 8 figg., 7 tavv.
Ili - BARBIERI F„ IACCARINO S.. BARBIERI F. & PETRUCCI F., 1967 - Il Pliocene del Subappennino Piacentino-Parmense-Reggiano. pp. 143-188, 20 figg., 3 tavv.
Giacomo Bracchi & Anna Alessandrello
Sezione di Paleontologia degli Invertebrati, Museo Civico di Storia Naturale di Milano
Paleodiversity of thè free-living polychaetes (Annelida, Polychaeta) and description of new taxa from thè Upper Cretaceous Lagerstàtten of Haqel, Hadjula and Al-Namoura (Lebanon)
Volume XXXII - Fascicolo III
Febbraio 2005
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
INDEX
Introduction . Pag. 3
Study area and geological setting . Pag. 5
Preservation . Pag. 5
Materials and methods . Pag. 6
Polychaete tossii record . Pag. 7
Systematic paleontology . Pag. 15
Paleobiology and paleoecology . Pag. 29
Discussion and conclusions . Pag. 30
Acknowledgements . Pag. 32
References . Pag. 33
Appendix . Pag. 40
© Società Italiana di Scienze Naturali
Museo Civico di Storia Naturale di Milano Corso Venezia ,55 -20121 Milano
In copertina: olotipo di Eunicites diopatroides n. sp. Foto di Roberto Appiani.
Registrato al Tribunale di Milano al n. 6694 Direttore responsabile : Anna Alessandrello Responsabile di redazione: Stefania Nosotti
Grafica editoriale: Michela Mura
Stampa: Litografia Solari, Peschiera Borromeo - Febbraio 2005
ISSN 0376-2726
Giacomo Bracchi & Anna Alessandrello
Paleodiversity of thè free-living polychaetes (Annelida, Polychaeta) and description of new taxa from thè Upper Cretaceous Lagerstàtten of Haqel, Hadjula and Al-Namoura (Lebanon)
Abstract - The present work reports thè results of a morphologic and taxonomic study of 377 specimens of free-living fossil polychates hold in thè collections of thè Museo Civico di Storia Naturale di Milano. The examined specimens come from thè Ceno- manian (Upper Cretaceous) Lagerstàtten of Haqel (361), Hadjula (15) and Al-Namoura (1) (Lebanon). A systematic definition at species level has been possible for 1 10 specimens from Haqel and 2 from Hadjula so as 6 families (Aphroditidae, Didonidae n. fam., Eunicidae, Goniadidae, Lumbrineridae and Oenonidae), 7 genera ( Didone n. gen., Eunicites, Ferragutia n. gen., Lumbriconerites, Paleoaphrodite, Phoeniciarabella n. gen. and Teruzzia n. gen.) and 17 species ( Didone pulcherrima n. gen. n. sp., Eunicites diopa- troides n. sp., Eunicites falcatus n. sp., Eunicites joinvillei n. sp., Eunicites mariacristinae n. sp., Ferragutia cenomaniana n. gen. n. sp., Lumbriconereites garassinoi n. sp., Lumbriconereites hadjulae n. sp., Paleoaphrodite libanotica n. sp., Phoeniciarabella caesaris n. gen. n. sp., Phoeniciarabella orensanzi n. gen. n. sp., Phoeniciarabella pinnulata n. gen. n. sp., Teruzzia gryphoeides n. gen. n. sp., Teruzzia pezzolii n. gen. n. sp., Teruzzia pusilla n. gen. n. sp. and Teruzzia sagittifera n. gen. n. sp.) of polychaetous annelids have been recorded from thè Cenomanian fossiliferous levels of Lebanon.
In addition to thè results of thè study of thè examined material is reported a systematic appraisal of thè fossil polychaetes where thè taxa described from Cambrian to Eocene are listed with regard to thè free-living forms.
Key words - Lebanon, Cenomanian, Polychaeta.
Riassunto - Paleodiversità dei policheti a vita libera (Annelida, Polychaeta) e descrizione di nuovi taxa dai giacimenti a conser¬ vazione eccezionale del Cretacico superiore di Haqel, Hadjula e Al-Namoura (Libano).
Sono riportati i risultati di un’indagine volta allo studio morfologico e tassonomico di 377 esemplari di policheti fossili conservati nelle collezioni del Museo Civico di Storia Naturale di Milano. I fossili esaminati sono riferibili a organismi a vita libera e provengono dai giacimenti a conservazione eccezionale di Haqel (361), Hadjula (15) e Al-Namoura (1) (Libano settentrionale), tutti di età ceno¬ maniana (Cretacico superiore). Gli esemplari per cui è stato possibile proporre una classificazione a livello di specie (1 12) provengono per la quasi totalità da Haqel (110), in minima parte da Hadjula (2). Tali esemplari sono stati attribuiti a 6 famiglie (Aphroditidae, Didonidae n. fam., Eunicidae, Goniadidae, Lumbrineridae e Oenonidae), 7 generi ( Didone n. gen., Eunicites, Ferragutia n. gen., Lum¬ briconerites, Paleoaphrodite, Phoeniciarabella n. gen. and Teruzzia n. gen.) e 17 specie ( Didone pulcherrima n. gen. n. sp., Eunicites diopatroides n. sp., Eunicites falcatus n. sp., Eunicites joinvillei n. sp., Eunicites mariacristinae n. sp., Ferragutia cenomaniana n. gen. n. sp., Lumbriconereites garassinoi n. sp., Lumbriconereites hadjulae n. sp., Paleoaphrodite libanotica n. sp., Phoeniciarabella caesaris n. gen. n. sp., Phoeniciarabella orensanzi n. gen. n. sp., Phoeniciarabella pinnulata n. gen. n. sp., Teruzzia gryphoeides n. gen. n. sp., Teruzzia pezzolii n. gen. n. sp., Teruzzia pusilla n. gen. n. sp. and Teruzzia sagittifera n. gen. n. sp.) eleggendo il giacimento di Haqel quale Lagerstàtten mesozoico per cui è nota la più ampia e varia fauna di organismi vermiformi.
Accanto all’esposizione dei risultati ottenuti dallo studio del materiale considerato viene riportata una check-list di famiglie, generi e specie di policheti fossili noti dal Cambiano all’Eocene, con riferimento alle sole forme a vita libera.
Parole chiave - Libano, Cenomaniano, Polychaeta.
INTRODUCTION
The first scientifìc treatise on thè fossil fauna from thè Upper Cretaceous Lagerstàtten of Lebanon was edited during thè first half of thè XIX century. The first paleontological note regarding thè Lebanese ichthyolithic limestones consists of a description of two new species of clupeomorph fish by De Blainville (1816) in “Nouveau Dictionnaire d’Histoire Naturelle”. The fossil fish from Lebanon had previously been cited by Herodotus (450 B.C.) and later by Sir Jean de Joinville (1248, 1309) in his memoirs about thè crusades of King Louis IX. The fossil ichthyofauna from thè Cretaceous Lagerstàtten in Lebanon
were also mentioned in various volumes published during thè XVIII century, such as thè following editions of thè “Voyage au Levant” (Le Brun, 1714; Banjer, 1725), “Travels or observations” (Shaw, 1738) and “Traité des Pétrifications” (Bourguet, 1742). The presence of both land plants and marine molluscs within thè same outcrops was first reported by Volney (1787). The toponym “Haqel” (also written as “Hakel” or “Haql”) first appears as “Heckel” in thè new edition of thè fifth volume of “Voyage au Levant” (Le Brun, 1725): nowadays this toponym reminds one of thè richest fossiliferous locality within thè Mesozoic.
4
GIACOMO BRACCHI & ANNA ALESSANDRELLO
There were many contributions on land plants, verte- brates and invertebrates from thè Lebanese Cretaceous Lagerstàtten in thè years following thè publication by De Blainville (1816). Agassiz (1833-1844), Egerton (1844), Heckel (1849), Pictet (1850), Pictet & Humbert (1866), Lewis ( 1 878) and Sauvage (1878) all increased thè knowl- edge about thè ichthyofauna from thè localities of Haqel and Sahel-Alma, describing several new genera and spe- cies of bony and cartilaginous fìsh. In thè following dec- ades new taxa of teleosts, sharks and rays were described by Dawis (1895), Woodward A.S. (1892, 1895a, 1895b, 1898, 1899a, 1899b, 1901, 1942), Hay (1903a, 1903b, 1903c), Roger (1946), Patterson (1967, 1970), Forey (1973, 1997), Capetta (1980), Goody (1969a, 1969b), Gaudant (1975, 1976, 1978a, 1978b), Gayet (1980a, 1980b, 1984, 1988), Tyler & Sorbini (1996), Bannikov & Bacchia (2000), Belouze & Gayet (2001), Brito & Dutheil (2001), Nursall & Capasso (2001), Zaragueta Bagils (2001), Poyato-Ariza & Wenz (2002), Capasso (2003) and Forey et al. (2003). Terrestrial vertebrates were first recorded by Lartet (1869) who referred to a fossil lizard from Sahel-Alma: thè actual systematic relationships of this reptile should be revised because Sahel-Alma is char- acterized by a particularly open sea facies (Dalla Vecchia, pers. comm.). More recently, Krassilov & Bacchia (2000) and Dalla Vecchia et al. (2002) mentioned thè exsistence of articulated phalanges assignable to a theropod dinosaur in a small store in Jbail, locality near Al-Namoura (also written as “en Nammoùra”, “Nammoura” or “Namoura”). The first avian skeletons from thè northem Gondwana has been also discovered in Al-Namoura together with some small feathers (Dalla Vecchia & Chiappe, 2002; Dalla Vecchia et al., 2002). Four tiny amber inclusions are exceptionally preserved in thè latter and are thè first record of fossil resin from Al-Namoura. However, thè presence of amber in fossiliferous outcrops in Lebanon had already been recorded from thè Lower Cretaceous localities of Hammana, Kdeirji, Mdeyrji and others, so as several nematodes, snails, mites, opilionids, spiders, insects, thè oldest reptile (lizard) in amber and thè first complete feather preseerved in a fossil resin have been described (Hennig, 1972; Schlee, 1973; Szadwieski, 1995, 1996; Prentice et al., 1996; Azar et al., 1999a, 1999b, 2000, 2003; Borkent, 2000; Grimaldi, 2001, 2004; Poinar & Milki, 2001; Podenas et al., 2001; Arnold et al., 2002; Azar & Nel, 2003; Penney & Selden, 2002; Lukaschevich & Mostovski, 2003; Penney, 2003; Perrichot et al., 2003; Nel et al., 2004). Moreover, Al Namoura has yielded new genera and species of semiaquatic (Dal Sasso & Pinna, 1997; Dal Sasso & Renesto, 1999) and flying reptiles (Dalla Vecchia et al., 2001) together with an hindlimbed snake (Rage & Escuillié, 2000, 2001, 2003; Rieppel & Head, 2004) and at least six specimens of chelonians (Dalla Vecchia et al., 2002). The notes by Russegger (1836), Roger (1946), Dilcher & Basson (1990), Dalla Vecchia & Venturini (1999), Krassilov & Bacchia (2000) and Dalla Vecchia et al., (2002) conceming thè algae and land plants (cycadophytes, auraucarias and angiospermae) from Hadjula (also written as “Hadjoula” or “Hjoula”) and Al-Namoura are also worth mentioning.
The first reference to Cretaceous invertebrates from thè Lebanese Lagerstàtten was by Kònig (1825) who illustrated two ophiuras from Haqel. Botta (1833) was thè first to mention thè crustacean fauna from Sahel-Alma
while Brocchi (1875) gave thè first scientific description of a fossil invertebrate from Haqel, thè decapod crusta¬ cean Penaeus libanensis. Fraas (1878) and Dames (1866) described and revised decapod crustaceans and cephalo- pods.
The decapod crustaceans from thè Upper Cretaceous Lagerstàtten in Lebanon represent thè most well-known aspect of thè invertebrate fauna discovered here to date. Thanks to thè works by Fraas (1878), Dames (1886), Van Straelen (1930, 1940), Glaessner (1945), Roger (1946), Forster (1984) and Garassino (1994, 2000, 2001), several specimens of macruran decapods have been assigned to thè Aegeridae, Benthesicymidae, Callianassidae, Erymidae, Glypheidae, Nephropidae, Oplophoridae, Palinuridae, Penaeidae, Scyllaridae and Stenopodidae. In addition to thè above, Garassino (1994) also described a new family, thè Carpopenaeidae. Withers (1928), Roger (1946) and Forster (1968) contributed to thè knowledge of thè brachyuran decapods from thè same Lagerstàtten : Forster (1968) assigned thè material he examined to thè Calappidae, whereas thè specimen formerly assigned to a living genus of Xanthiidae by Roger (1946) has been recently revised as a new genus of thè Necrocarcinidae (Larghi, 2004). Other Cretaceous brachyurans from Leba¬ non have been assigned to a new species of Raninidae (Dames, 1866; Glaessner, 1945; Roger, 1946), even if thè type material needs revision (Garassino, pers. comm.), and to a new subfamily and a new genus of Dorippidae (Larghi & Garassino, 2000; Larghi, 2004). Dames (1886), Van Straelen (1938), Roger (1946), Teruzzi (1983), Schram et al. (1999) and Lange et al. (2000) have also reported thè presence of cirriped, mysidacean, stomato- pod and thylacocephalan crustaceans.
Apart from thè crustacean studies, few investiga- tions of thè other invertebrates preserved in thè Upper Cretaceous Lagerstàtten of Lebanon have been car- ried out and thè fauna is mainly stili unrecorded. Some data have been reported by Fraas (1878), Woodward H. (1879, 1883), Kolbe (1888), A.S. Woodward (1896), Naef (1922), Roger (1946), Engeser & Reitner (1986), Forey et al. (2003), Novati (2003) and Fuchs (in press) regarding foramminifers, molluscs, horseshoe crabs, insects, opilionids and echinoderms. This is probably a reflection of thè methods applied in thè collection of thè fossils: thè Lebanese limestones have been and are often excavated by thè locai population that disregard small fossils of no aesthetic interest, because of their limited economical value. Consequently, very little data about Lebanese fossil polychaetes have been collected to date by Roger (1946) and Kozur (1971). The former author studied some mouth apparata, scolecodonts and incom¬ plete specimens from Haqel and Hadjula. He assigned a maxillary apparatus and an incomplete specimen to thè fossil eunicid Eunicites Ehlers, 1868 and an incomplete specimen to thè living phyllodocid Phyllodoce Savigny, 1818. Two poorly preserved specimens were described by Roger (1946) as indeterminable “worms” or “annelids?”. In thè same way, Kozur (1971) described two polychaete specimens from Haqel as indeterminate eunicids. How¬ ever, in recent years, due to an increasing interest in some previously neglected invertebrate taxa, several specimens of cephalopods, nematodes, polychaetes and echiurans from thè Lebanese Lagerstàtten are now housed within thè collections of some scientific institutions, such as thè
PALEOD1VERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
5
Museo Civico di Storia Naturale di Milano (Alessandrello et al., 1995). To date this has resulted in two publications regarding descriptions of a new polychaete species, Eunicites phoenicius Alessandrello & Teruzzi, 1986, and a new tetrameric nematode (Alessandrello & Mattavelli, in press), both from Haqel.
Aims of thè present study are thè description of approximately 400 specimens of free-living polychaetes from thè Cenomanian of Lebanon as well as a system- atic appraisal, with regard to thè free-living forms, of thè fossil polychaetes where thè taxa described from Cam- brian to Eocene are listed.
STUDY AREA AND GEOLOGICAL SETTING
The examined specimens come from thè sub-litho- graphical limestones of Haqel, Hadjula and Al-Namoura, thè most celebrated fossiliferous localities of Lebanon, renowned world-wide for thè variety and thè excellent conservation of thè preserved fauna. These localities are located in north-west Lebanon, some kilometres north- east of Beirut, together with thè other paleontological sites of Sahel-Alma, Maifouk and Dahr-el-Kelb. The sites of Haqel and Hadjula are positioned 12 km from thè Lebanese coast, at an altitude of approximately 700 m above sea level. They are about 4 km distant from each other and 45 km from Beirut. Maifouk and Dahr-el-Kelb are located a few kilometres north of Hadjula, whereas thè 15 m to 30 m thick limestones of Al-Namoura out- crop closer to thè coast, 25 km south of Haqel and 20 km north of Beirut.
Geological studies on thè Cretaceous outcrops of Lebanon were started in thè second half of thè XIX cen- tury by Botta (1833), who investigated thè lithology of thè Haqel and Sahel-Alma areas. Some years later, Heckel (1849) assigned an Upper Cretaceous age to Sahel-Alma, while Pictet & Humbert (1866) suggested a slightly older age for Haqel. The fìrst geological data conceming thè locality of Hadjula was published by Lewis (1878) and Fraas (1878), who considered both Haqel and Hadjula as Turonian in age. However, thè Cenomanian age of Haqel, Hadjula and Maifouk was later recognized by Douvillé (1910), Zumoffen (1926) and Patterson (1967). Blankehom (1926), Dubertert & Vautrin (1937) and Ejel & Dubertret (1966) dated Sahel-Alma back to thè Senon- ian. The biostratigraphical studies of Dubertret (1959,
1966), Hiickel (1969, 1970, 1974a, 1974b) and Saint Marc (1974) placed thè sequences at Haqel, Hadjula and Maifouk at thè Lower-Middle Cenomanian bound- ary. According to Hemleben (1977) Haqel and Hadjula are younger: he dated back these sites to thè late Ceno¬ manian. Hiickel (1970) also attested thè younger age of thè Haqel strata that actually occur twenty metres higher stratigraphically than those of Hadjula. At thè same time, thè strata at Sahel-Alma and Al-Namoura have been dated back, through thè investigation of thè foraminifera fauna, to thè late Senonian (Ejel & Dubertret, 1966) and to thè middle Cenomanian (Dalla Vecchia & Venturini, 1999; Dalla Vecchia et al., 2002; Forey et al., 2003) respec- tively.
According to Hiickel (1969, 1970, 1974a, 1974b), Hemleben (1977) and Capetta (1980), thè sub-litho- graphical limestones from Haqel and Maifouk are hard, fine grained, well-bedded and laminated, often charac- terized by a rich fossiliferous content, a yellowish colour that sometimes may become greyish. Some levels of thè Haqel outcrop are particularly rich in flint nodules. The same authors describe thè limestone from Hadjula as a more compact, soft and laminated rock, characterized by a lighter yellow or grey-yellow colour and without flint nodules. In comparison with thè rocks from thè other Lebanese sites, thè limestone from Sahel-Alma is lighter coloured (Capetta, 1980). The limestone from Al-Namoura is very thinly laminated, green to grey-blue in colour and is rich in flint nodules (Dalla Vecchia & Venturini, 1999).
PRESERVATION
The favourable conditions present for fossilization during thè late Cretaceous in Lebanon are expressed in thè state of preservation of thè polychaetes and other soft-bodied metazoans. Compared with those from other Lagerstàtten thè polychaetes from Haqel are among thè best preserved fossil worms known (Briggs & Kear, 1993). With regard to thè lightly skeletized tis- sues, thè fossil polychaetes from Lebanon show preser¬ vation of thè cuticle (samples MSNM Ì26329 (Fig. 1), MSNM Ì26335 and MSNM Ì26338), resistant against degradation thanks to its structure of collagen fibers arranged in crossed strata and enriched with biopoly- mers (Storch, 1988; Stankiewicz et al., 1997), thè chae- tae, in living polychaetes formed of sclerotinized chitin and inorganic material (Specht, 1988; Butterfield, 1990a), and thè pharyngeal apparatus. The latter varies greatly in composition in thè modern polychaetes (Olive, 1980) but it is generally formed of sclerotinized
collagen mineralized with calcite or aragonite (Voss- Foucart^a/., 1973; Fauchald & Rouse, 1997). In some taxa, such as thè oenonids and certain glyceroideans, thè jaws and maxillae are strenghtened with metal- ions, such as iron, copper or zinc (Colbath, 1986, 1988; Colbath & Larson, 1987; Purschke, 1988; Eriksson & Elfman, 2000), and with organic compounds, such as tanned proteins, aromatic aminoacids, glycerine and hystidine (Bergman, 1989). In thè material examined here, thè chaetae and pharyngeal apparata are preserved both as molds and mineralized structures but thè latter is more common than thè former. In some specimens thè mineralized elements that comprise thè pharyngeal apparatus are black and shiny, in others they are whit- ish and pearly. The former type of fossilization often involves also thè three-dimensionally preserved cha¬ etae. In thè examined specimens thè lighter coloured material is resulted typical of thè mouth apparatus of
6
GIACOMO BRACCHI & ANNA ALESSANDRELLO
thè labidognath eunicidans, such as thè eunicids and lumbrinerids, while thè darker material is resulted typi- cal of thè prionognath eunicidans, such as thè oenonids (Colbath, 1986; Fauchald & Rouse, 1997). Some of thè specimens examined (MSNM Ì9355 for example) possess a pharyngeal apparatus impregnated by a very flaky and whitish minerai. With regard to soft-tissue preservation, thè fossil polychaetes from Lebanon generally show traces of thè alimentary canal and body outline only, although two anal cirri have been recognized by thè authors in a lumbrinerid specimen (MSNM il 3301) and in an undetermined specimen
(MSNM Ì23414). Some longitudinal muscle bundles are evident in two goniadid samples (MSNM il 2423 and MSNM Ì24961). Worthy of note are thè traces of pigmentation in thè undetermined specimen MSNM Ì26326 (Fig. 2), a feature already observed by Arduini et al. (1982) in a fossil lumbrinerid from thè prealpine Lagerstàtte at Osteno (Lower Jurassic), northern Italy. Most of thè samples examined in thè present study have usually been found flattened along thè bedding piane and they show a dorso-ventral compression, even if in a few cases a lateral compression is evident.
Fig. 1 - MSNM Ì26329, undetermined specimen preserving thè cuti- cle.
Fig. 2 - MSNM Ì26326, undetermined specimen preserving traces of pigmentation.
MATERIALS AND METHODS
There are 379 polychaete specimens (see Appendix) from thè Upper Cretaceous of Lebanon housed in thè paleontological collections of thè Museo Civico di Storia Naturale di Milano (MSNM). Two specimens from Haqel have already been described by Alessandrello & Teruzzi (1986) as Eunicites phoenicius. Among thè samples examined here, 362 are from Haqel, 15 from Hadjula (MSNM 25107, MSNM Ì25108, MSNM i25 109, MSNM Ì25110, MSNM Ì25113, MSNM Ì25115, MSNM Ì25116,
MSNM Ì25119, MSNM Ì25120, MSNM Ì25125, MSNM Ì25443, MSNM Ì26332, MSNM Ì26335, MSNM Ì26336 and MSNM Ì26338) and 1 (MSNM Ì9363) from Al- Namoura.
The systematic relationships of thè examined speci¬ mens have been recognized, where possible, through comparisons with both fossil and living species.
The polychaete taxonomy adopted in this work essen- tially follows Fauchald (1977).
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
7
POLYCHAETE FOSSIL RECORD
The polychaete fossil record includes traces, articu- lated pharyngeal apparata, isolated elements of thè jaw apparatus and, more rarely, fully preserved specimens (Howell, 1962; Hàntzschel, 1975; Wills, 1993). The isolated elements of thè pharyngeal apparata of thè euniceans and glyceroideans are particularly common in Paleozoic deposits: they have been defined by Croneis & Scott (1933) as “scolecodonts”. Since thè erection of a taxonomic scheme based on fragmentary specimens does not comply with thè rules of thè International Code of Zoological Nomenclature (Aldridge, 1990) and in con- sideration of thè thè diffìculty of comparing articulated apparata, whole-preserved specimens and scolecodonts, it follows that thè latter should have, in thè same way as thè conodonts, a biostratigraphical and paleoecological value but not a useful systematical signifìcance (Zaw- idzka, 1975; Bergman, 1989, 1995, 1998; Hints, 1999a; Eriksson, 1998; Eriksson & Bergman, 2003; Eriksson & Leslie, 2003). This is for example thè case of thè arabellid genus Synclinophora Eisenack, 1975 from thè Silurian of Sweden and Poland, recently re-described by Mierzejew- ski (1984) based on an articulated maxillary apparatus. This apparatus is difficult to compare with those of thè arabellid specimens from Lebanon that we have described here. Even though thè Lebanese arabellids also have thè jaw apparatus and part of thè body preserved, however, in most cases, thè maxillae are lacking. Thus, for a long time, most of those studying thè polychaetes have argued that parataxonomical investigations of thè scolecodonts should be kept separate from thè orthotaxonomical stud- ies on full-preserved specimens (Eller, 1964; Bengston, 1985a; Tasch & Stude, 1965). A parataxon has been defined by Melville (1959) as “a taxon based on a frag- ment [...] which can be classifìed at thè genus-group and species-group levels [...] but cannot be assigned to thè same (ortho)taxa at those levels as thè whole animai to which they belong”. However, some authors, such as Snajdr (1951), Kielan-Jaworovska (1961, 1968) and Kozur (1970, 1971), have tried to combine thè orthotaxo¬ nomical and parataxonomical investigations of thè fossil polychaetes. Consequently, over thè last 40-50 years, some authors have used thè scolecodonts to reconstruct articulated apparata (Lange, 1949; Kozlowski, 1956; Sylvester, 1959; Kielan-Jaworovska, 1962, 1966; Sza- niawski, 1968, 1970, 1974; Kozur, 1971; Szaniawski & Wrona, 1973; Corradini & Olivieri, 1974; Jansonius & Craig, 1974, 1975; Mierzejewski & Mierzejewska, 1975; Mierzejewski, 1978; Szaniawski & Gadzicki, 1978; Berg¬ man, 1979, 1981; Mànnil & Zaslavskaya, 1985a, 1985b; Colbath, 1987; Szaniawski & Imajima, 1996; Hints, 1999a, 1999b; Eriksson & Bergman, 1998; Eriksson, 2003a). This method has been criticized by Jansonius & Craig (1971), Szaniawski & Wrona ( 1 973) and Thompson & Johnson (1977), although according to Bergman (1989, 1991) some Paleozoic scolecodonts should be orthotaxo- nomically investigated and compared both at thè genus and species level. On thè one hand, thè application of this method has resulted in thè resolution of some problems within polychaete systematics, thanks to thè recognition of several “form genera” (Edgar, 1984), or rather in thè recognition of some scolecodonts as belonging to later discovered jaw apparata or whole-preserved specimens
(Eriksson & Bergman, 1998; Nakrem et al, 2001). On thè other hand, thè same method has contributed to confu- sion within polychaete systematics, both because a high number of synonyms have been yielded when scoleco¬ donts occur in more than one form genus and as some parataxa have been absorbed by some orthotaxa. In even worse cases, some scolecodonts have been assigned to previously described orthotaxa , thus generating serious problems in thè use of thè same orthotaxa during thè description of newly discovered fully preserved speci¬ mens. This is, for instance, thè case of Eunicites Ehlers, 1 868 and Lumbriconereites Ehlers, 1 869, both based on full-preserved specimens but later adopted in order to classify several scolecodonts.
As pointed out by Howell (1962) and Wills (1993) polychaete traces, scolecodonts, jaw apparata and full- preserved specimens have been recorded in Paleozoic, Mesozoic and Cenozoic rocks, ranging from thè Cam- brian to thè Pleistocene. The presence of polychaete annelids within thè Precambrian stili requires defini¬ tive confirmation (Xiao et al, 2001; Sepkoski, 2002). Several Ediacaran fossils have been assigned to thè Polychaeta (Glaessner, 1976a, 1976b, 1979) however, even if in some cases they appear to actually possess some polychaete characters, their systematic relation- ships remain unsubstantiated. Only Spriggina Glaessner, 1958 has been definitively accepted as an arthropod- like metazoan. The genera Parmia Gnilovskaya, 1998, Pararenicola, Wang, 1983 Ruedemannella Howell, 1969 and Protoarenicola Wang, 1983 from thè Upper Riphean (Lower Algonkian) of Russia (Timan biota) and China (Huainan biota) are considered to be non-free-living “annelidomorphs” (Gnilovskaya et al., 2000), enigmatic polychaetes (Howell, 1965) or enigmatic metazoans (Sun et al., 1986; Chen, 1988). However, some authors (Qian et al., 1999) have recently re-interpreted Protoarenicola and Ruedemannella, thè latter found also in thè Pridolian (Upper Silurian) of New York State (Howell, 1959), as a megascopic alga, while other researchers (Sun et al., 1986; Chen, 1988; Xiao et al. 2001) suspect thai Pararen¬ icola and Protoarenicola are sponges.
In thè following paragraphs an annotated check-list has been compiled for thè free-living forms of thè fossil polychaetes ranging in age from Cambrian to Eocene. Only those with a description based on full-preserved specimens have been included. Families and genera are ordered alphabetically. The fact that thè fossil free-living polychaetes need a generai systematic revision clearly emerges from thè check-list, in particular with regard to thè Mesozoic euniceans as well as thè taxa described in thè first half of thè XX century or earlier. Moreover, thè check-list underlines that thè fossil record of thè whole- preserved free-living polychaetes is rather poor due to thè essentially soft-bodied nature of these invertebrates: taking into account also thè taxa of essentially uncer- tain affinities, approximately only 1 5-20% of thè extant families have been recognized within thè fossil fauna. Full-preserved polychaetes are more abundant where exceptional conditions of fossilization prevail: in addition to that of Haqel, some famous Lagerstdtten such as thè Burgess Shale (Cambrian, British Columbia), Bear Gulch (Carboniferous, Montana), Mazon Creek (Carbonifer-
8
GIACOMO BRACCHI & ANNA ALESSANDRELLO
ous, Illinois), Grès à Voltzia (Triassic, France), Osteno (Jurassic, Italy), La Voulte sur-Rhòne (Jurassic, France), Solnhofen (Jurassic, Bavaria), Ponte Giurino (Triassic, Italy) and Bolca (Eocene, Italy), have yielded several examples of exceptionally preserved specimens (Briggs & Clarkson, 1987; Alessandrello, 1990a; Briggs & Kear, 1991; Etter, 2002a, 2002b; Hagadom, 2002a, 2002b; Schellenberg, 2002; Tang, 2002; Alessandrello & Brac¬ chi, 2004; Alessandrello et al., 2004).
An updated polychaete fossil record including both parataxa and sedentary forms is given in Sepkoski (2002).
Family Amphinomidae Savigny in Lamarck, 1818
(Carboniferous-Recent)
Full-preserved amphinomids have been recorded from thè Upper Carboniferous of Europe and North America, more precisely from thè Kasimovian-Gzelian (Upper Car¬ boniferous) of Montceau-les-Mines (Pacaud et al., 1982; Rolfe et al., 1982; Heyler, 1986; Heyler & Poplin, 1988), France, and from Mazon Creek (Meek & Worthen, 1865; Thompson, 1979). The only Mesozoic full-preserved amphinomids have been described by Ehlers (1869) from thè Solnhofen Plattenkalk. The specimens described by Ehlers (1869) and Thompson (1979) retain imprints of thè soft body as well as bunches of bristly spines. In thè living organism, these bristles were probably filled with poison. Some modem amphinomids possess nearly identical poi- sonous spines which justify thè nickname “fìreworms” used for these polychaetes.
Meringosoma Ehlers, 1869 (Jurassic; Germany) - This genus includes only thè type species M. curtum Ehlers, 1869, characterized by a short (2-3 cm), ovai and flattened body equipped with bristles at thè mid- length. However, in thè opinion of thè authors and in consideration of what can be deduced from thè avail- able plates and photos, thè systematic relationships of Meringosoma should be revised in respect of thè recent discovery of several aphroditids in some Mesozoic Lagerstàtten.
Paleocampa Meek & Worthen, 1865 (Carboniferous; Illinois and France) - Meek & Worthen (1865) described a full-preserved worm-like organism from thè Coal Meas- ures of Illinois as P. anthrax Meek & Worthen, 1865. Thompson (1979) later described about thirty amphino- mid specimens from thè famous “Pit 11” mine of Mazon Creek as Raphidiophorus hystrix Thompson, 1979. In thè following years several authors (Pacaud et al., 1982; Rolfe et al., 1982; Heyler, 1986; Heyler & Poplin, 1988; Briggs & Kear, 1993) reported full-preserved polycha¬ etes referable to Paleocampa from thè Carboniferous of Montceau-les-Mines, suggesting that Paleocampa and Rhaphidiophorus are synonyms. According to Schellen¬ berg (2002), thè taxonomic relationships of Paleocampa within thè polychaetes is uncertain: a further revision of thè fossil amphinomids is needed in order to resolve this issue and to verify thè relationships between Paleocampa and Raphidiophorus. The latter was described by Thomp¬ son (1979) as an epifaunal or pelagic, probably camivo- rous amphinomid hearing tentacular cirri, an unarmed pharynx and biramous parapodia with heavy and protec- tive chaetae. The body of R. hystrix is short (20-25 mm) and includes few segments.
Superfamily Aphroditacea Fauchald, 1977
(Devonian?-Recent)
In thè most recent linnean treatment of thè poly¬ chaete taxonomy (Fauchald, 1977), thè superfamily Aphroditacea includes six families often indicated as “scale-worms”: Acoetidae Kingberg, 1856, Aphroditidae Malmgren, 1867, Eulephetidae Chamberlin, 1919, Poly- noidae Malmgren, 1867, Pholoidae Kingberg, 1858 and Sigalionidae Malmgren, 1867. In thè aphroditacean fos- sils thè recognition of thè characters that can be useful to hypothesize affinities with one of these families is rather problematic, particularly in thè distinction among aphro¬ ditids, pholoids, polynoids and sigalionids. Since only a few species have been assigned doubtfully to thè Pholoi¬ dae, Polynoidae and Sigalionidae in thè aphroditacean fossil record, thè definition of thè scale-worms as a super¬ family is considered by thè authors of thè present work as being more suitable to thè usuai situations observed for thè fossils, in which reai affinities with thè aphroditacean families are, in most cases, only suspect.
Fossils which could be tentatively assigned to Sigalio¬ nidae have been recorded in thè Ordovician of Minnesota (U.S.A.) by Stauffer (1933), who described thè parataxon Thalenessites Stauffer, 1933, on thè basis of some poly- gonous scales and jaws. More recently Alessandrello et al. (2004) have described thè enigmatic aphroditacean Protopholoe Alessandrello, Bracchi & Riou, 2004 from La Voulte-sur-Rhòne and pointed out some similarities between this Jurassic genus and thè modem sigalionid Pholoe Johnston, 1839.
Some affinities with thè Polynoidae have been pointed out for both Paleozoic and Mesozoic aphro- ditaceans: these are thè Carboniferous scale-worms from Mazon Creek (Thompson, 1979) and thè Triassic scale-worms from Grès a Voltzia (Gali & Grauvogel, 1966).
The fossil record of full-preserved specimens which could be assigned to Aphroditidae includes both Paleo¬ zoic (Clarke, 1903; Thompson, 1979) and Mesozoic (Alessandrello & Teruzzi, 1986b; Alessandrello, 1990b; Alessandrello et al., 2004) genera. The Cenozoic genus Sthenelaites Rovereto, 1904 was proposed in order to absorbe (Rovereto, 1904) thè Eocenic species Nere- ites dasiaeformis Massalongo, 1855 and it has been included among thè aphroditids (Howell, 1962; Sepkoski, 2002). However, Alessandrello (1990a) recently revised Sthenelaites and identified it as a seaweed of thè family Dasycladaceae. Thompson (1979) included thè Carbon¬ iferous aphroditids from Mazon Creek within thè new subfamily Hystriciolinae Thompson, 1979. The hystricio- line aphroditids seem to be thè most common polycha¬ etes from Mazon Creek: they are jawed aphroditids that possess an eversible proboscis and biramous parapodia equipped with at least five types of chaetae that are how¬ ever, lacking thè elytrae. Other whole-preserved aphro¬ ditids have been recorded from both Gondwanan and Laurasian Lagerstàtten of Mesozoic age (Alessandrello & Teruzzi, 1986b; Alessandrello et al., 2004). These Tri¬ assic and Jurassic aphroditids show evidence of both jaws and elytrae occurring together with biramous parapodia and a variety of setal types. The fossil aphroditids are in need of systematic revision, in particular regarding thè Mesozoic taxa.
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCR1PTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
9
Dryptoscolex Thompson, 1979 (Carboniferous; Illi¬ nois) - Hystricioline aphroditid known only on thè basis of thè type species D. matthiesae Thompson, 1979 from Mazon Creek. Dryptoscolex was probably an epifaunal polychaete, in life habits very similar to thè non-commen- sal species of thè modem family Polynoidae Malmgren, 1867 (Thompson, 1979). D. matthiesae is one of thè few fossil aphroditids to possess an elongate worm-like body and a pharynx armoured with some jaw pieces.
Fastuoscolex Thompson, 1979 (Carboniferous; Illi¬ nois) - Predaceous aphroditids assigned by Thompson (1979) to thè subfamily Hystriciolinae but strictly related to thè epifaunal aphroditids. The genus includes only thè type species F. gemmatus Thompson, 1979 from Mazon Creek: it is one of thè few fossil aphroditids to possess a pharyngeal armature (Thompson, 1979).
Homaphrodite Gali & Grauvogel, 1966 (Triassic; France) - The type species Homaphrodite speciosa Gali & Grauvogel, 1966 from Grès à Voltzia is thè only species recognized within this genus. The actual systematic rela- tionship of Homaphrodite within thè polychaetes should be revised through comparisons with thè other Mesozoic aphroditids. This is particularly necessary as Thompson (1979) argued that H. speciosa should probably be moved to thè Polynoidae Malmgren, 1867 because it possesses a pair of dorsoventral maxillae whose structure is quite similar to that of thè modem representatives of thè family (Gal & Grauvogel, 1966). However, a similar pattern of thè pharyngeal armature has already been observed in both living (Fauchald & Rouse, 1997) and fossil aphro¬ ditids (Alessandrello et al, 2004). In thè opinion of Gali & Grauvogel (1966) Homaphrodite possesses elytrae.
Hystriciola Thompson, 1979 (Carboniferous; Illinois) - Epifaunal and free-living aphroditid known only on thè basis of thè type species H. delicatula Thompson, 1979 from Mazon Creek. It is thè most common aphroditid in thè Coal Measures of Illinois (Thompson, 1979).
Paleoaphrodite Alessandrello & Teruzzi, 1986 (Tri- assic-Cretaceous; Italy, France, Madagascar and Leba- non) - Most of thè Mesozoic aphroditids found to date are included within this genus, which seems strictly related to thè extant genus Aphrodita Linnaeus, 1758. This is especially true with regard to thè generai morphology and shape of thè body, which is typically similar to that of thè sea-mice, as well as for thè architecture of thè parapodial chaetae. In addition to thè type species P. raetica Ales¬ sandrello & Teruzzi, 1986 from thè Triassic of thè Italian Prealps, thè only species of thè group that seems to pre¬ serve thè elytrae (Alessandrello & Teruzzi, 1986b), thè following have been recorded to date: P. anaboranoensis Alessandrello, 1990 from thè Olenekian (Lower Triassic) of Madagascar, P. ade/iae Alessandrello, Bracchi & Riou, 2004, P. briggsiana Alessandrello, Bracchi & Riou, 2004 and P gallica Alessandrello, Bracchi & Riou, 2004 from La Voulte sur-Rhòne, France. P briggsiana and P. adeliae are thè largest fossil aphroditids known (10-15 cm) and they are very similar to each other in thè generai mor¬ phology of thè body. However, these two Jurassic species can be distinguished on thè basis of thè architecture of thè jaw pieces that armour thè pharynx: they are thè only species of thè genus that preserve these structures (Ales¬ sandrello et al., 2004). P. briggsiana also shows evidence of thè existence of an eversible pahrynx within thè genus, while P adeliae preserves a felt of capillary fibers on thè
dorsum. As they are naturally unlikely to be fossilized, capillary fibers must be considered, in thè same way as thè elytrae, as a poor diagnostic feature in systematic studies of Paleoaphrodite (Alessandrello & Teruzzi, 1986a; Alessandrello et al., 2004). A new species P. liba- notica n. sp. from Haqel is described in thè present study. The gut content retained by one specimen of P libanotica suggests a predatory habit for this genus, since it includes some fish vertebrae. The six species included within Pale¬ oaphrodite to date should be revised by comparison with thè other Mesozoic aphroditids.
Protonympha Clarke, 1903 (Devonian; Canada) - The type species P. salicifolia Clarke, 1903 is thè oldest aphroditid known. This poorly known taxon has been found in thè Devonian shales of Ontario County, Naples, Canada.
Family Burgessochaetidae Conway Morris, 1979
(Cambrian)
Monospecifìc family established by Conway Morris (1979) subsequent to thè revision of some specimens from thè Burgess Shale formerly assigned to Canadia Walcott, 1911. The relationships of thè family Burgesso¬ chaetidae within thè class Polychaeta are uncertain ( Wills, 1993). However, among thè modem polychaetes, Amphi- nomidae and Nephtyidae seem to show dose similarities with thè notosetae and neurosetae of thè bugessochaetids (Conway Morris, 1979). According to Fauchald & Rouse (1997) only some phyllodocid traits can be inferred from thè fossils of Burgessochaeta.
Burgessochaeta Conway Morris, 1979 (Cambrian; Canada) - The only species known within this genus is thè type species B. setigera (Walcott, 1911). Burgessochaeta is interpreted as a burrower that propelled itself on cha¬ etae (Conway Morris, 1979).
Family Canadiidae Walcott, 1911
(Cambrian)
This monospecifìc family seems to share some simi¬ larities with thè modem chrysopetalids (Conway Morris, 1979; Butterfield, 1990a) but its relationship within thè class Polychaeta are considered uncertain (Wills, 1993). Together with thè wiwaxiids, thè canadiids have been included into thè fossil superfamily Canadiacea Butter¬ field, 1 990 within Phyllodocida. According to Fauchald & Rouse ( 1 997) only some phyllodocid traits can be inferred from thè fossils of Canadia. Selkirkia Walcott, 1911 from thè Burgess Shale had also been previously assigned to thè canadiid polychaetes (Howell, 1962) but it is now assumed to be a priapulid (Wills & Sepkoski, 1993).
Canadia Walcott, 1911 (Cambrian; Canada) - The type species C. spinosa Walcott, 1911 is thè only spe¬ cies that is stili included within this genus after Conway Morris (1979) revised thè Burgess Shale polychaetes. The other taxa referred to Canadia by Walcott (1911) have been split between thè burgessochaetids and peronocha- etids by Conway Morris (1979). The same author inter¬ preted Canadia as a benthic and free-living polychaete. According to Butterfield (1990a) C. spinosa must prob¬ ably be affiliated to thè chrysopetalids.
10
GIACOMO BRACCHI & ANNA ALESSANDRELLO
Family Chrysopetalidae Ehlers, 1864
(Carboniferous?- Recent)
While investigating thè polychaete fauna from Mazon Creek, Thompson (1979) referred to thè presence of undescribed polychaetes of thè family Palmyridae King- berg, 1858. Palmyrids are nowadays included among thè chrysopetalids, with thè exception of Palmyra Savigny, 1818, which has been moved to thè aphroditids (Fauchald & Rouse, 1997). The Mazon Creek “palmyrids” represent thè only fossils known within thè Chrysopetalidae, even if Conway Morris (1979, 1985) and Butterfield (1990a) have pointed out that Canadia spinosa and Wiwaxia corrugata from thè Burgess Shale show some modem chrysopetalid-like traits.
Family Didonidae n. fam.
(Cretaceous)
The institution of this family is based on thè descrip- tion, proposed in this paper, of five specimens of large- bodied eunicemorph polychaetes from Haqel. The didonids show intermediate characters between thè labi- dognath eunicidans and thè dorvilleids. Furthermore, thè most significant feature of thè didonids is thè maxillary apparatus equipped with an unpaired carrier. Among thè modem and free-living eunicidans such a carrier can be found in some dorvilleids. However, since thè carrier of thè five specimens from Flaqel is not strongly linked to thè forceps, we prefer to relate them to a new group of thè oenonid-dorvilleid line.
Bidone n. gen. (Cretaceous; Lebanon) - The type spe- cies D. pulcherrima n. gen. n. sp. from Haqel is newly described in thè present study.
Family Eunicidae Berthold, 1827
(Ordovician-Recent)
The oldest full-preserved fossil eunicids are Carbonif¬ erous in age, but thè family also includes some Paleozoic scolecodonts from Europe and North America (Howell, 1962).
Esconites Thompson & Johnson, 1977 (Carbonifer¬ ous; Illinois) - This genus includes only thè type species E. zelus Thompson & Johnson, 1977 from Mazon Creek. The systematic relationships of Esconites within thè Eunicidae need to be revised. According to Thompson & Johnson (1977) these full-preserved eunicids from Mazon Creek cannot be assigned to Eunicites, even if their mor- phology appears to be strictly related with it, as numerous scolecodonts have previously been assigned to this taxon. However, thè originai description of Eunicites edited by Ehlers (1868) was based on a whole-preserved specimen and not on isolated jaws or maxillae. Thus, thè full-pre¬ served specimens from Mazon Creek could eventually be related to thè ort ho taxon Eunicites, while thè scoleco¬ donts should eventually be assigned to other parataxa. In thè same way as Eunicites , also Esconites appears to be strictly related to thè extant genus Eunice Cuvier, 1817.
Eunicites Ehlers, 1868 (Jurassic-Eocene; Germany, Italy and Lebanon) - Some full-preserved specimens assigned to Eunicites have been found within thè Meso¬
zoici E. triasicus Gali & Grauvogel, 1966 from Grès à Voltzia and E. orobicus Alessandrello & Bracchi, 2004 from thè Italian Prealps are Triassic in age, E. proavus (Germar, 1 842) from Solnhofen is Jurassic in age while E. phoenicius Alessandrello & Teruzzi, 1986 from Haqel is Cretaceous in age. Ehlers (1868) described thè type species E. avitus Ehlers, 1868 based on thè description of thè genus Eunicites for some eunicid polychaetes from Solnhofen. Ehlers ( 1 869) later described E. atavus Ehlers, 1 869 and E. dentatus Ehlers, 1 869 from thè same outcrop and proposed thè binomial E. proavus as a new combination for Geophilus proavus Germar, 1842. Some years later Kozur (1971) stated that E. avitus, E. atavus, E. dentatus and E. proavus are synonyms of E. avitus : following thè rules of thè International Code of Zoologi- cal Nomenclature, E. proavus has priority. However, it is considered here that by closely studying thè plates of thè manuscripts on thè worms from thè Solnhofen Plat- tenkalk (Ehlers, 1868, 1869) it can be clearly deduced that thè synonymity proposed by Kozur (1971) is rather doubtful: further studies are required to confimi this issue. While revising thè polychaetes from Monte Bolca (Massalongo, 1850, 1855), Alessandrello (1990a) also described E. affinis (Massalongo, 1855), E. gazolae (Massalongo, 1855), E. intermedia (Massalongo, 1855) and E. pinnai Alessandrello, 1990. All thè other species described under Eunicites (Hinde, 1879, 1880, 1882, 1896; Foerste, 1888; Patte, 1921; Eller, 1934a, 1934b, 1938, 1940, 1941, 1942, 1944, 1945, 1955, 1963a, 1964; Seidel, 1959; Sylvester, 1959; Taugourdeau, 1968, 1970) were not based on full-preserved specimens but on scole¬ codonts. These scolecodonts are hardly comparable with thè full-preserved species of thè same genus or with thè genus description fumished by Ehlers (1868) in particu- lar. Consequently, according to thè mles of thè I.C.Z.N., thè classification of these scolecodonts should be kept separated from thè orthotaxon Eunicites by thè institu¬ tion of a different parataxon or parataxa. In this paper new specimens of full-preserved Eunicites from Haqel are described as E. joinvillei n. sp., E. falcatus n. sp., E. diopatroides n. sp. and E. mariacristinae n. sp. The most significant features for distinguishing thè full-preserved species assigned to date to thè Eunicites reside in thè pha- ryngeal armature. The shape of thè jaws (subtriangular, scythe-shaped or subelliptical), thè pattern of thè anterior borders (smooth or denticulate) and thè shape of thè car- riers, which are usually paired, triangular, joined along a median line or sometimes equipped with small lateral and rounded paddles {E. diopatroides n. sp.), seem to be particularly diagnostic.
Family Fossundecimidae Thompson, 1979
(Carboniferous)
Thompson (1979) instituted this family to include thè second most common polychaete species from Mazon Creek. According to Thompson (1979), thè fossundecim- ids seems phyllodocids that show intermediate characters between thè hystricioline aphroditids and nereidids. They have a similar paleoecology, extemal morphology and architecture of jaws and chaetae as thè former and thè number and architecture of thè jaws is thè same as in thè latter.
PALEODIVERSITY OF THE FREE-L1VING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
11
Fossundecima Thompson, 1979 (Carboniferous; Illi¬ nois) - Epifaunal and probably omnivorous polychaetes known on thè basis of thè type species F. konecniorum Thompson, 1979 only. Thompson (1979) hypothesized that F. konecniorum could be a primitive ancestor of thè nereidid polychaetes.
Family Goniadidae Kingberg, 1865
(Carboniferous-Recent)
The goniadids have been included within Glyceridae for a long time (Fauchald & Rouse, 1997). Full-preserved goniadids have been found to date within thè Paleozoic, more precisely in thè Carboniferous strata ofNorth Amer¬ ica (Schram, 1 979; Thompson, 1 979). In thè present paper thè Mesozoic Ferragutia n. gen. from Haqel is described. Moreover, thè goniadid fossil record includes several Triassic scolecodonts (macrognaths) formerly assigned to thè parataxon Alienites Kozur, 1970 but later moved to thè extant genus Goniada Adouin & Milne Edwards, 1833. However, Szaniawski (1974) and Zawidzka (1975) believe that thè assigment of these isolated macrognaths to a living genus is rather dubious as thè architecture of thè macrognaths among thè goniadids is very similar. Therefore, these structures do not represent a diagnostic character in thè systematic definition of these polychaetes. Moreover, Goniada is not a parataxon : this implies that it cannot be adopted in order to classify isolated macrog¬ naths or other isolated and disjunct glyceroidean jaws.
Carboserostris Schram, 1979 (Carboniferous; Mon¬ tana) - The type species C. megaliphagon Schram, 1979 is thè only one known within this genus. It is thè most abun- dant species among thè polychaetes from Bear Gulch. The stomodaeum of Carboserostris is equipped with microg- naths and macrognaths. The latter have not been observed in Ferragutia n. gen. and Pieckonia (see below).
Ferragutia n. gen. (Cretaceous; Lebanon) - The genus is established here based on thè description of thè type species F. cenomaniana n. gen. n. sp., thè second most common polychaete among those from Haqel. The sto¬ modaeum of Ferragutia n. gen. is equipped with both micrognaths and chevrons. The latter have not been observed in Carboserostris and Pieckonia (see above and below).
Pieckonia Thompson, 1979 (Carboniferous; Illinois) - This genus includes only thè type species P. helenae Thompson, 1979. It is considered to be a rapid burrower, in thè same way as thè modem goniadids. The stomo¬ daeum of Pieckonia is equipped with micrognaths how¬ ever, differently from Carboserostris and Ferragutia n. gen. (see above), it lacks both macrognaths and chevrons. Ferragutia n. gen. is lacking thè macrognaths as does Pieckonia , a condition that Thompson (1979) explains by hypothesizing a primitive condition.
Family Hesionidae Grube, 1850
(Carboniferous-Recent)
Only a dozen polychaete specimens from Europe and North America have been assigned to thè hesionids to date: two specimens from Monte Bolca (Alessandrello, 1990b), ten specimens from Mazon Creek (Thompson, 1979).
Rutellifrons Thompson, 1979 (Carboniferous; Illi¬ nois) - At present this genus includes only thè type spe¬ cies R. wolfforum Thompson, 1979 from Mazon Creek. The specimens described by Thompson ( 1 979) are worthy of note due to thè preservation of various soft parts, such as thè eversible muscular proboscis together with thè ten- tacular and parapodial cirri. R. wolfforum was probably an epifaunal polychaete.
Siphonostomites Rovereto, 1904 (Eocene; Italy) - Rovereto (1904) described under this genus thè material reported by Massalongo (1855) as Nereites hesionoides Massalongo, 1855. Alessandrello (1990b) confìrmed thè revision of Rovereto (1904) and pointed out a strong similarity between Siphonostomites and thè extant genus Hesione Savigny, 1818.
Family Insolicoryphidae Conway Morris, 1979
(Cambrian)
Monospecific family set up by Conway Morris ( 1 979), who pointed out that insolicoryphids do not show charac- ters that could suggest dose similarities with any modem polychaete or with thè other monospecific families from thè Burgess Shale.
Insolicorypha Conway Morris, 1979 (Cambrian; Canada) - The type species I. psygma Conway Morris, 1979 has been interpreted as a pelagic swimmer.
Family Lumbrineridae Schmarda, 1861
(Ordovician-Recent)
To date full-preserved lumbrinerids have been recorded from both Mesozoic (Ehlers, 1869; Arduini et al., 1982) and Paleozoic (Schram, 1979) strata and have been assigned to three genera. Moreover, Howell (1962) has documented thè presence of thè living genus Lumbrineris Grube, 1840 from thè Pliocene to recent times. The new Cretaceous genus Teruzzia n. gen. is newly described in this study. According to Szaniawski & Imajima (1996) thè lumbrinerids originated from some representati ves of thè family Hartmaniellidae Imajima, 1977, such as Lysaretides Kozur, 1970, during thè Upper Cretaceous. Infact, thè hartmaniellids now include both Lysaretides and Delositesl Kozur, 1 967, genera formerly included within thè family Lysaretidae Kingberg, 1865 emend. Kozur, 1970 but later included within thè Lum¬ brineridae and Oenonidae respectively (Colbath; 1989; Orensanz, 1990; Fauchald & Rouse, 1997). However, thè Cretaceous specimens described in thè present work and thè Carboniferous Phiops are very similar to thè modem representatives of thè family and do not show any kind of primitive character. Thus, it is considered here that thè origin of thè lumbrinerids can probably be dated back to thè Middle-Upper Paleozoic by thè systematic revi¬ sion of various scolecodonts erroneously assigned to thè orthotaxon Lumbriconereites (Hinde, 1879, 1882; Foer- ste, 1888; Stauffer, 1933, 1939; Eller, 1938, 1941, 1942, 1945, 1946, 1963a, 1964, 1969; Taugourdeau, 1968; Seidel, 1959; Sylvester, 1959; Jansonius & Craig, 1971; Tasch & Stude, 1966). The “ Lumbriconereites scoleco¬ donts” should therefore be revised systematically with thè eventual institution of new parataxa.
12
GIACOMO BRACCHI & ANNA ALESSANDRELLO
Lumbriconereites Ehlers, 1869 (Ordovician?-Cre- taceous; Germany) - Genus described on thè basis of full-preserved specimens assigned to thè type species
L. deperditus Ehlers, 1869, that must consequently be considered an orthotaxon. However, as mentioned above, some polychaete researchers assigned several scoleco- donts from Paleozoic and Mesozoic rocks to Lumbricon¬ ereites and thus adopted thè genus like a parataxon. Since thè originai description of Lumbriconereites proposed by Ehlers (1869) is not based on scolecodonts but on whole- preserved specimens, it follows that thè scolecodonts for- merly assigned to this genus should probably be moved to one or more new parataxa. However, thè reai systematic value of Lumbriconereites has been questioned by Kozur (1971), who argued that L. deperditus is a synonym of Eunicites proavus, another eunicemorph polychaete from Solnhofen. It is considered in thè present study, that by observing thè plates of thè manuscript on thè worms from thè Solnhofen Pìattenkaìk (Ehlers, 1869), one can clearly deduce that there are differences in thè architecture of thè jaws of E. proavus and L. deperditus : further studies are required to confimi this issue. In thè present work thè new species L. hadju/ae n. sp. and L. garassinoi n. sp. from thè Haqel and Hadjula localities are described.
Melanoraphia Arduini, Pinna & Teruzzi 1982 (Jurassic; Italy) - Genus known only on thè basis of thè type species M. maculata Arduini, Pinna & Temzzi 1982, from thè Italian Prealps. The most interesting feature of
M. maculata is thè presence of pigmentation traces on thè parapodia (Arduini et al., 1982). In thè opinion of thè authors thè relationships of Melanoraphia within thè Eunicida need to be revised as thè absence of thè antennae in thè available specimens, used as a decisive character in thè recognition of thè systematic affinities of thè genus, could be an artifact of fossilization. Furthermore, thè pha- ryngeal armature of M. maculata is that of a typical labi- dognath eunicidan and its generai architecture together with thè presence of little semicircular paddles on thè sides of thè carriers as well as thè absence of antennae seem to suggest eunicid-like traits.
P/iìops Schram, 1979 (Carboniferous; Montana) - The specimens from thè Carboniferous of Bear Gulch described under thè binomial P aciculorum Schram, 1979 are noteworthy for thè good state of preservation of thè lobate parapodia. The genus can be clearly distin- guished from thè other fossil lumbrinerids simply on thè basis of thè subtriangular shape of thè jaws which, differ- ently from that of Lumbriconereites and Teruzzia n. gen., shows no similarity with thè species included in thè living Lumbrineris.
Teruzzia n. gen. (Cretaceous; Lebanon) - In thè same way as Lumbriconereites , Teruzzia n. gen. appears to be strictly related to thè living genus Lumbrineris. Teruzzia n. gen. includes, in addition to thè type species T. pezzolii n. gen. n. sp., T. gryphoeides n. gen. n. sp., T. sagittifera n. gen. n. sp. and T. pusilla n. gen. n. sp., and represents thè most common polychaete taxon in thè Haqel outcrop.
Family Nephtyidae Grube, 1850
(Carboniferous-Recent)
Full-preserved polychaetes of this family are known only from thè Paleozoic of North America and they have
been assigned to two genera (Thompson, 1979; Schram, 1979). With approximately one thousand specimens discovered to date, thè nephtyids seem to be thè most common free-living polychaetes from Mazon Creek, even if in some cases their systematic affinities are questioned.
Astreptoscolex Thompson, 1979 (Carboniferous; Illinois and Montana) - The type species A. anasillosus Thompson, 1979, from thè Coal Measures of Mazon Creek (Thompson, 1979) and from Bear Gulch (Schram, 1979) was probably gregarious. Thompson (1979) inter- preted Astreptoscolex as an epifaunal creeper.
Didontogaster Thompson, 1979 (Carboniferous; Illi¬ nois) - The type species D. cordylina Thompson, 1979, with approximately 650 specimens found to date, is thè second most common polychaete from Mazon Creek. Recently Sutton et al. (200 la) have considered thè rela¬ tionships of Didontogaster with thè nephtyids as doubtful, even if thè characters pointed out by Thompson (1979) seem to confimi these affinities with reasonable certainty.
Family Nereididae Johnston, 1865
(Ordovician-Recent)
Since thè Eocenic species described by Massalongo (1855) under thè living genus Nereis Linnaeus, 1758 have been included within thè eunicid genus Eunicites (Alessandrello, 1990a), full-preserved fossil nereidids are recorded only from Mesozoic and Paleozoic strata. However, thè nereidid affinities of thè three fossil genera listed below have to be newly demonstrated as they are recognized on thè basis of old notes. Thus, thè data available regarding thè full-preserved fossil nereidids should be updated by an adequate revision that takes into account thè current scientifìc developments in fossil polychaete research: meanwhile, their inclusion among thè nereidids is considered provisional. With regard to thè Paleozoic, several parataxa have been related to this family (Hinde, 1879, 1880; Stauffer, 1933, 1939;Zebera, 1935; Eisenack, 1939; Eller, 1940, 1941, 1942, 1945, 1955, 1963a, 1963b, 1964, 1969; Sylvester, 1959; Tasch & Stude, 1966; Kozur, 1967; Taugoudeau, 1968, 1970; Jansonius & Craig, 1971). The Lower Cambrian Fauciv- ermis yunnanicus Hou & Chen, 1989 has been tentatively assigned to thè nereidid polychaetes (Hou & Chen, 1989; Chen & Erdtmann, 1991; Delle Cave & Simonetta, 1991) although, in thè opinion of Fauchald & Rouse (1997), thè presence of polychaetes in thè Lower Cambrian cannot be confirmed to date. This is due to thè fact that thè Lower Cambrian worm-like fossils from China seem to not show any characters that can definitively demonstrate a nereidid or polychaete affìnity.
Ctenoscolex Ehlers, 1869 (Jurassic; Germany) - This genus includes only thè type species C. procerus Ehlers, 1 869 from Solnhofen. The specimens described by Ehlers (1869) as polychaetes of doubtful but probable nerei¬ did affinities, were re-examined by Kozur (1970) who argued that Ctenoscolex is not a polychaete as it lacks thè chaetae in spite of an excellent preservation. The same author (Kozur, 1971) later suggested that Ctenoscolex be considered as related to an uncertain family within thè Phyllodocida.
Nawnites Roy, 1929 (Devonian; New York) - Poorly known taxon from thè Devonian of North America. It
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
13
has been described on thè basis of thè type species N. gilboensis Roy, 1929 only. Roy (1929) fumished few morphological characters of this polychaete and limited his description to thè segmented body.
Triadonereis Mayer, 1954 (Triassic; Germany) - Genus strictly related to thè living genus Nereis and described on thè basis of thè type species T. eckertii Mayer, 1954 from thè Trochitenkalk (Middle Triassic) of Germany. The suggested relationships of Triadonereis link it to thè neridids (Mayer, 1954) but its actual affinities within thè polychaetes must be demonstrated.
Family Oenonidae Kingberg 1865
(Silurian?-Recent)
This family presently includes all thè genera previ- ously included within Arabellidae Hartman, 1944 and some genera formerly assigned to Lysaretidae (Colbath, 1989; Orensanz, 1990; Fauchald & Rouse, 1997). To date thè fossil record of thè oenonid polychaetes comprises some Paleozoic scolecodonts and articulated apparata from thè Silurian of thè Baltic area and North America that have been assigned to Synclinophora Eisenack, 1975 (Mierzejewski, 1984; Eriksson, 2003b) as well as several Mesozoic scolecodonts from centrai Europe that have been described by Kozur (1970, 1971) under thè modem genera Arabella Grube, 1850, Halla Costa, 1844, Drilo- nereis Clarapède, 1870, Notocirrus Schmarda, 1861 and Oenone Savigny, 1818. However, Szaniawski (1974) and Zawidzka (1975) have already expressed doubts regarding thè assignment of these Mesozoic scolecodonts to modem oenonid genera with any reasonable certainty. Moreover, thè genera mentioned by Kozur (1970, 1971) are ortho- taxa that cannot be considered within thè parataxonomi- cal investigation of thè scolecodonts. In thè present study thè fìrst full-preserved oenonid genus within thè Meso¬ zoic has been described: it includes three species. In thè opinion of Kielan-Jaworovska (1966), Szaniawski (1970) and Szaniawski & Gadzicki (1978), thè oenonids arose from thè parataxonomical family Atraktoprionidae during thè Middle-Upper Paleozoic, even if Mierzejewski (1984) argued that oenonids and atraktoprionids are sister taxa that share a common ancestor.
Phoeniciarabella n. gen. (Cretaceous; Lebanon) - Large-bodied oenonids strictly related to thè living genus Arabella. Among thè examined material, apart from thè type species P. pinnulata n. gen. n. sp., P. caesaris n. gen. n. sp. and P orensanzi n. gen. n. sp. were also recognized. Phoeniciarabella n. gen. represents thè only oenonid genus known within thè Mesozoic. All thè species of thè genus possess elongate and slender carriers in thè same way as thè modem oenonids, but they seem to lack thè unpaired piece that is usually associated to thè maxillary carriers of thè latter.
Family Peronochaetidae Conway Morris, 1979
(Cambrian)
Monospecific family established by Conway Morris (1979) after revison of specimens from thè Burgess Shale formerly assigned (Walcott, 1911) to Canadia. The relationships of thè family Insolicoryphidae within thè
Polychaeta are uncertain (Wills, 1993). However, in thè opinion of Conway Morris (1979) thè uniramous parapo- dia of thè peronochaetids recali thè posterior neuropodia of thè modem Paraonidae Cerniti, 1909.
Peronochaeta Conway Morris, 1979 (Cambrian; Canada) - Conway Morris (1979) interpreted thè type species P. dubia Walcott, 1911 as a burrower and a scav- enger.
Family Phyllodocidae Òrsted, 1843
(Devonian-Recent)
Together with those of thè nereidids and sigalionids, thè phyllodocid fossil record is one of thè most reduced among thè full-preserved free-living polychaetes. To date whole-preserved phyllodocids have not been recorded within thè Mesozoic (Howell, 1962), apart from one specimen from thè Upper Cretaceous of Sahel-Alma, Lebanon, doubtfully assigned by Roger (1946) to thè living genus Phyllodoce Savigny, 1818. The earliest records of thè fossil phyllodocids are from thè Middle- Late Paleozoic of North America where thè majority of thè specimens assigned to thè family have been found.
Levisettius Thompson, 1979 (Carboniferous; Illinois)
- Genus tentatively assigned to thè phyllodocids based on thè description of thè only known species L. campy- lonectus Thompson, 1979, interpreted as a free-living and camivorous polychaete (Thompson, 1979). However, in thè opinion of thè authors, thè inclusion of Levisettius within thè phyllodocids should be considered provisionai and as doubtful as thè specimens described by Thompson (1979) lack thè decisive traits that could demonstrate a phyllodocid affinity.
Palaeochaeta Clarke, 1903 (Devonian; New York)
- Clarke (1903) assigned this genus to thè aphroditids and described only thè type species P devonica Clarke, 1903. However, thè same author indicated thè resemblance between Paleochaeta and thè modem representatives of Phyllodoce and Nereis, intrinsically anticipating its assign¬ ment to thè phyllodocids (Howell, 1962; Sepkoski, 2002).
Family Stephenoscolecidae Conway Morris, 1979
(Cambrian)
Monospecific family established by Conway Morris (1979), who pointed out only a superficial similarity with thè modem Iospiliidae Bergstròm, 1914.
Stephenoscole x Conway Morris, 1979 (Cambrian; Canada and Utah) - The type species S. argutus Conway Morris, 1979 is thè worst preserved polychaete among those from thè Burgess Shale. Conway Morris tentatively suggested that this species was benthic. According to Briggs & Kear (1991) also some specimens from thè Middle Cambrian Spence Shale of Utah can be doubtfully assigned to Stephenoscolex.
Family Tomopteridae Johnston, 1865
(Carboniferous-Recent)
Probably due to thè pelagic mode of life and thè absence of jaws typical of these polychaetes, their fossil
14
GIACOMO BRACCHI & ANNA ALESSANDRHLLO
record is reduced to a few Paleozoic specimens (Briggs & Clarkson, 1987). However, according to Glaessner (1976a), also Spriggina floundersi Glaessner, 1958 from thè Vendian of thè Ediacara Hills, Australia, can be con- sidered a tomopterid polychaete. Wills & Sepkoski (1993) included Spriggina within thè enigmatic class Paratrilo- bita Fedonkin, 1985 which comprises some problematic Precambrian taxa that have been variously assigned to thè Arthropoda and Annelida. More recently McMenamin (2003) defined Spriggina as a trilobitoid ecdysozoan and underlined its arthropod affìnites.
Eotomopteris Briggs & Clarkson, 1987 - (Carbonif- erous; Scotland) - Genus based on thè description of thè type species E. aldrigei Briggs & Clarkson, 1987, from thè Lower Carboniferous of thè Granton Shrimp Bed, near Edinburgh (Scotland). The exceptional preservation of these Paleozoic tomopterids allows evidence of ten- tacular cirri and parapodia to be observed.
Family Wiwaxiidae Walcott, 1911
(Cambrian)
The systematic relationships of this monospecifìc family from thè Burgess Shale have been discussed for a long time. Even if Wills & Sepkoski (1993) treat thè wiwaxiids as problematic metazoans, according to But- terfield (1990a) they show some affinities with thè Phyl- lodocida. However, together with Canadia, they should be included in thè tossii superfamily Canadiacea Butterfield, 1990 which shares some characters with both thè Chrysop- etalidae and Aphroditidae. More recently, Conway Morris & Peel (1995) and Conway Morris & Gould (1998) under¬ lined that thè wiwaxiids are closely related to thè halkieri- ids, problematic invertebrates from thè Lower Cambrian (Bengston & Missarzhevsky, 1981; Jell, 1981; Bengston & Conway Morris, 1984; Bengston, 1985b; Conway Morris & Peel, 1990), because both groups have very similar sclerites. The same authors pointed out that thè halkieri- ids show some primitive characters that link them to thè polychaetes and brachiopods and that thè wiwaxiids might be a sister taxon to thè Polychaeta. This view seem to be accepted also by Dzik (2003) who put thè halkieriids along thè same lineage of thè brachiopods, molluscs and poly¬ chaetes even if thè transitional stages among these groups have not yet been found in thè fossil record. According to Butterfield ( 1 990a) Wiwaxia and Canadia are more closely related to each other than either are to any other fossil organism and that sclerites such as theirs have been widely adopted by disparate groups of organisms.
Wiwaxia Walcott, 1911 (Cambrian; Canada and Utah) - The type species W corrugata (Matthew, 1899) was first described by Matthew (1899), thanks to a single spine found across thè valley of thè Burgess Shale, where full-preserved specimens were later discovered (Walcott, 1911). Wiwaxia was interpreted as a primitive mollusc or an elytigerous scale-worm for a long time (Walcott, 1911; Butterfield, 1990a). However, Conway Morris (1985) and Butterfield (1990a) investigated thè microstuctural details of some sclerites and noticed that they are very similar to thè paleae (flattened chaetae) of some modem chrysopeta- lids and aphroditids. According to Briggs & Kear (1991) also some specimens from thè Middle Cambrian Spence Shale of Utah can be tentatively assigned to Wiwaxia. In
thè opinion of Conway Morris & Gould (1998) Wiwaxia might be a member of thè annelid stem group, or rather a creature stili in thè process of becoming an annelid. In thè opinion of Fauchald & Rouse (1997) thè systematic posi- tion of Wiwaxia has yet to be clarified even if it clearly shows some chrysopetalid-like traits.
Unrelated and problematic genera
Acaenoplax Sutton, Briggs, Siveter & Siveter, 2001 (Silurian; England) - Vermiformi organism from thè Her- efordshire Lagerstàtte of England, originally described as an aplacophoran mollusc (Sutton et ai, 200 lb, 200 le, 2004). However, this systematic interpretation has been argued by Steiner & Salvini-Plawen (2001) who proposed to consider A. hayae Sutton, Briggs, Siveter & Siveter, 2001 as an highly specialized polychaete that shares some characters with thè modem Aphroditidae, Chaetopteridae, Cirratulidae and Stemaspidae.
Eopolychaetus Ruedemann, 1901 (Ordovician; New York) - Only thè type species E. albaniensis Ruedemann, 1901, from thè Canajoharie Shale of New York State, has been assigned to this genus. According to Wills (1993) and Sepkoski (2002), Eopolychaetus belongs to an unnamed family within an uncertain order.
Kenostrychus Sutton, Briggs, Siveter & Siveter, 2001 (Silurian; England) - On thè basis of a cladistic analysis, K. clementsi Sutton, Briggs, Siveter & Siveter, 2001, from thè Silurian volcanoclastic deposit of Her- efordshire, has been interpreted as a member of a stem group from which both thè Phyllodocida and Eunicida originate (Briggs et al., 1996; Sutton et al., 200 la).
Myoscolex Glaessner, 1979 (Cambrian; Australia) - The systematic relationships of M. ateles Glaessner, 1979 from thè Lower Cambrian of Kangaroo Island (South Australia) have been discussed for a long time (Conway Morris, 1979; Glaessner, 1979; Wills, 1993; Briggs & Nedin, 1997; Dzik, 2003). In thè opinion of Glaessner (1979) thè anatomy of Myoscolex link it to thè echiurids but according to Conway Morris (1979) and Briggs & Nedin (1997) both echiurid and polychaete affinities are questionable. Recently Dzik (2003) re-discussed thè ana- tomical features of Myoscolex and proposed to consider it as one of thè oldest polychaete annelid known, super- ficially similar to thè modem opheliids. Moreover, Dzik (2003) pointed out thè resemblance between Mysocolex and thè Cambrian Pikaia gracilens Walcott, 1911 from thè Burgess Shale which has been originally described as an annelid (Walcott, 1911) but later interpreted as a chor- date (Conway Morris, 1979).
Pontobdellopsis Ruedemann, 1901 (Ordovician; New York) - This genus includes only thè type species P. cometa Ruedemann, 1901, a poorly known taxon from thè Ordovician of thè Canajoharie Shale, New York State. According to Wills (1993), this genus belongs to an unnamed family within an uncertain order.
Protopholoe Alessandrello, Bracchi & Riou, 2004 (Jurassic; France) - Enigmatic aphroditacean recently ; described from La Voulte sur-Rhòne on thè basis of a single pyritized specimen. Alessandrello et al. (2004) pointed out that Protopholoe shares some characters with thè living Pholoe, a genus actually included among thè sigalionid aphroditaceans.
PALEODfYERSITY OF THE FREE-LIVING POLYCHAETES (ANNELID.V POLYCHAETA i AND DESCRIPTION OF NEW TAXA EROSI THE EPPER CRETACEOL'S OF LEBANON
15
Ramesses Schram, 1979 (Carboniferous: Montana)
- The type species R. magnus Schram, 1979 is charac- terized by parapodia similar in some respects (Schram. 1979) to those of thè modem Orbiniidae Hartman, 1942. According to Wills (1993) and Sepkoski (2002) this genus belongs to an unnamed family within an uncertain order. However. thè modem orbiniids are not free-living forms as Ramesses seems to be.
Soris Schram. 1979 (Carboniferous; Montana)
- Schram (1979) indicated thè type species S. labiosus Schram. 1979 as belonging to an undetermined poly- chaete order. The same author pointed out, through thè
investigation of thè pharyngeal armature of thè only t\vo available specimens. a superficial similarity with some Paleozoic parataxa such as Glycerites blinde, 1879 and Paranereites Eisenack. 1939. According to Wills (1993) and Sepkoski (2002) this genus belongs to an unnamed family within an uncertain order.
Trentonia Pickerill & Forbes, 1978 (Ordovician: Canada) - The type species is T. shegiriana Pickerill & Forbes, 1978 from thè Ordovician of thè Quebec City area (Trenton Series). To date Trentonia has been doubtfully considered an enigmatic eunicemorph polychaete (Pick¬ erill & Forbes, 1978; Wills, 1993; Sepkoski, 2002).
SYSTEMATIC PALEONTOLOGY
As clearly emerges from thè check-list compiled above and as already pointed out by Fauchald & Rouse (1997), thè systematic paleontology of thè free-living polychaetes can be very confusing. even to those studying thè poly¬ chaetes. This is essentially due to thè soft-bodied nature of these invertebrates. Thus, several diagnostic characters related to soft parts and adopted in thè systematic neontol- ogy of thè polychaetes, such as thè cirri, thè antennae, thè gills. thè parapodia or thè elytrae, are usually not evident in thè fossils. This is also thè case even when thè latter are represented by whole-preserved specimens.
Most of thè characters adopted in thè systematic paleontology of thè polychaetes derive from thè jaw pieces that armour thè pharynx in some groups such as thè eunicidans. glyceroideans. some aphroditaceans. thè nephtyids and others, and from other structures resist- ant to degradation. such as thè chaetae. even if thè latter usually do not presene their diagnostic ultrastructure. This situation implies that thè fossil taxa can hardly be incorporated within thè linnean schemes of thè polycha¬ ete taxonomy. Such a problem is. for example, evident in thè paleontological investigation of thè labidognath euni¬ cidans and aphroditaceans. Among thè former thè preser- vation of thè lateral and median antennae can be useful in distinguishing thè eunicids from thè lumbrinerids. despite a very similar phaiyngeal armature (George & Hartman- Schròder. 1985; Fauchald & Rouse, 1997). In thè fossil eunicids that show full-presen ation. thè presen ation of thè antennae has, for example. only been documented in one specimen of thè Cretaceous species Eunicites phoeni- cins from Febanon (Alessandrello & Teruzzi. 1986a). However, in thè other species of thè same genus these soft structures have never been observed. even if all thè euni¬ cids should possess them by definition. On thè contrary. thè living lumbrinerids are lacking antennae (George & Hartman-Schròder. 1985: Fauchald & Rouse. 1997) while in thè Jurassic eunicidan species Melanoraphia maculata. thè absence of antennae has been considered a decisive factor in thè assignment of this taxon to thè lumbrinerids (Arduini et al., 1982), despite thè fact that thè species has a very similar pharyngeal armature, especially in thè architecture of thè jaws and carriers, to that of certain Mesozoic eunicids. However. thè soft-bodied nature of thè antennae make thè possible preservation of these prostomial structures very unlikely in fossils: according to this overview thè absence of antennae in Melanoraphia and in most of thè species assigned to thè Eunicites could therefore be an artifact of fossilization. Consequently. thè
affinities of these taxa cannot be considered as proven at all if thè rules of thè systematic neontology of thè polychaetes are followed. With regard to thè aphrodita¬ ceans. thè elytrae and thè dorsal felt of capillary fibers are diagnostic soft-bodied features which are very rarely preserved in thè fossil forms: for example, thè Triassic species Paleopahrodite raetica and thè Jurassic species Paleoaphrodite adeliae respectively have thè elvirae and thè dorsal felt preserved. However, these structures have not been observed in thè other species of thè same genus. Thus, thè elytrae and dorsal felt of thè aphroditaceans are. differently from what is outlined in thè systematic neontology of thè group (Day, 1967; Fauchald & Rouse. 1997), not useful as diagnostic features for their definition within systematic paleontology.
With thè exception of those from thè Middle Cambrian of thè Burgess Shale. mostly represented by carbonaceous remains ( Butterfield. 1990b), most of thè fossil polycha¬ etes are too poorly preserv ed to be characterized using thè rules of systematic neontology. The information that can be deduced from these fossils is limited and an actual systematic. taxonomic and phvlogenetic analysis is sub- stantially not possible even if some authors (Orensanz. 1990) have tried to include some fossil taxa within their systematic treatment of thè modem polychaetes. Thus. thè systematic paleontology of thè free-living polychaetes must be based on characters that are often evident in thè fossil forms but that have little significance in a system¬ atic review of thè modem polychaetes. A similar situation is found with regard to thè systematic paleontology of other worm-like invertebrates. such as thè sipunculans and enteropneusts (Cutler. 1994; Alessandrello et al.. 2004).
Phylum Annelida Lamarck. 1 809 Class Polychaeta Grube. 1850 Order Eunicida Dales. 1962 Family Eunicidae Savigny, 1818 Genus Eunicites Ehlers. 1 868
Eunicites joinxillei n. sp.
PI. I
Diagnosis: subelliptical jaws not protruding beyond thè lateral borders of thè forceps. closely positioned side by side along a centrai junction line and disposed at an angle of almost 180°. Jaws with denticulate front borders
16
GIACOMO BRACCHI & ANNA ALESSANDRELLO
equipped with 4 teeth, strongly rounded lateral borders and slightly convex back margins. Separated, triangular, elongate and longitudinally grooved mandibular proc- esses. Short, triangular and symmetrical carriers joined along a centrai junction line.
Derivatio nominìs : dedicated to Sir Jean de Joinville, a crusader of King Louis IX, who first mentioned (1305- 1308) thè existence of thè Lebanese fossils.
Geologica l age : Cenomanian.
Type locality : Haqel.
Material : 5 specimens. MSNM il 2420 (holotype), MSNM il 24 14 and MSNM il 3304 (paratypes), MSNM Ì20612, MSNM Ì25456.
Description : elongate and thin body measuring from 140 mm (MSNM Ì12420) to 290 mm (MSNM Ì20612) in length and from 2.5 mm (MSNM il 24 14) to 9 mm (MSNM i 1 2420) in width. The body width is Constant with thè exceptions of MSNM Ì20612, in which it decreases from thè cephalic to thè caudal extremity, and of MSNM Ì25456, in which it slightly increases at thè level of thè pharyngeal region. The measurements of thè examined specimens are typical of a large bodied polychaete.
Only thè stomodaeal apparatus and a short (30 mm) part of thè cephalic extremity are preserved in MSNM i 1 3304 and MSNM Ì25456, while MSNM Ì12414 (220 mm) lacks thè caudal extremity. The other specimens are full-preserved.
In MSNM il 3304 thè jaw apparatus is preserved as an imprint, while mineralized jaws and molds of thè maxil- lary apparatus are present in thè other specimens. How- ever, small, mineralized fragments of thè anterior part of thè latter are evident in MSNM il 2420, MSNM Ì20612 and MSNM Ì25456. The pharyngeal armature measures from 4.2 mm (MSNM Ì25456) to 13 mm (MSNM il 2420) in length. In thè holotype thè jaws measure 3 mm in length and 1.2 mm in width.
The anterior ends of arcuate and hooked forceps are always preserved but only in MSNM il 24 14 (3.8 mm) and MSNM Ì25456 (1.7 mm) is thè distance between them measurable. Excluding MSNM il 24 14, in which a small fragment of thè left forcep is mineralized, thè for¬ ceps are always preserved as imprints.
The carriers are always poorly preserved as imprints with thè exception of MSNM Ì20612, in which two small, mineralized fragments (15-20 mm) are evident along thè centrai junction line.
The forceps and carriers are only not preserved in MSNM i 1 3304.
The poor quality of preservation of thè examined specimens does not allow thè total number of metameres to be counted even if, on thè basis of thè aciculae arrange¬ ment, at least 300 metameres (0.8 mm in thickness) can be counted in MSNM Ì20612, 170 (0.5 mm in thickness) in MSNM il 24 14, 100 (1 mm in thickness) in MSNM il 2420 and 50 in MSNM Ì25456.
Three aciculae equip each parapodium. Each parapo- dium bears two elongate (0.8 mm in MSNM Ì25456, 3 mm in MSNM il 24 14) and thickened aciculae toward thè prostomium and a third shorter and thinner acicula that appear to be divaricated from thè longer ones and tumed backwards. So as in thè living Eunice, thè slenderest acicula could be interpreted as notopodial, thè thickest as neuropodial (Fauchald, 1992). In MSNM il 24 14 thè length of thè two longest aciculae decreases from thè
cephalic extremity (3 mm) to thè caudal one (2 mm) and in MSNM Ì25456 thè parapodia bear tufts of two parallel and thick aciculae that show thè same length. However, in consideration of thè stomodaeal apparatus architecture of MSNM il 24 14 and MSNM Ì25456, which is very similar to that of thè other specimens that are assigned here to E. joinvillei n. sp., thè aciculae arrangement must be consid- ered as being poorly diagnostic due to thè intraspecific variability and thè fossilization artifacts. The position in which thè notopodial and neuropodial aciculae of MSNM Ì25456 and MSNM il 24 14 are preserved sug- gests that thè ventral surface is being observed in these specimens. Probably thè same specimens were subjected to compression during diagenesis so that, in thè fossils, thè notopodial acicula appear to be located behind thè neuropodial acicula. Morover, in thè same specimens, thè jaw apparatus seems to be superimposed on thè maxillary apparatus: since in thè eunicemorph polychaetes thè jaws are located along thè ventral side of thè pharynx, this kind of preservation confirms that it is thè ventral surface of these specimens that is being observed. On thè contrary, thè position in which thè elements of thè pharyngeal apparatus of MSNM il 2420 are preserved suggests that thè dorsal surface is being observed.
Traces of thè digestive tube are preserved as dark- brown traces in MSNM il 24 14, MSNM il 2420 and MSNM Ì25456.
Remarks : thè complex architecture of thè pharyngeal armature of thè examined specimens clearly marks them as eunicidan polychaetes. Moreover, thè pattern of thè jaws, thè shape of thè paired carriers and thè kind of preservation of thè pharyngeal apparatus, sometimes con- sisting of a white and pearly material quite similar to thè originai one, suggest that they belong to thè labidognath eunicids or lumbrinerids (Fauchald, 1992; Fauchald & Rouse, 1997). It is preferred here to assign thè material examined to Eunicites, especially due to thè strong simi- larity of thè stomodaeal armature (Ehlers, 1868, 1869), even if thè eunicids should possess, in contrast to thè lum¬ brinerids, prostomial antennae that were, however, not observed in thè specimens (George & Hartman-Schròder, 1985). Among thè full-preserved Eunicites described to date, grooved mandibular processes very similar to those of E. joinvillei n. sp. have already been recorded (Ehlers, 1868; Kozur, 1970; Alessandrello, 1990) in E. proavus and E. affinis. Furthermore, thè jaws of E. proavus are subelliptical like those of E. joinvillei n. sp., but they are not denticulate or sharpened at thè lateral ends. Moreover, in E. proavus thè jaws are separated by an angle of about 120-140°, as well as having totally joined and smooth surfaced mandibular processes: on thè contrary in E. join¬ villei n. sp. thè same angle measures almost 1 80° and thè mandibular processes show a grooved surface. The jaws of E. affinis have denticulate front borders like those of E. joinvillei n. sp. but they show a regular ovai shape and slightly sharpened lateral ends. Moreover, thè mandibular processes of E. affinis are grooved like those of E. join¬ villei n. sp., but they are joined for 2/3 of their length, differently from thè completely separated mandibular processes of E. joinvillei n. sp.
In addition to E. joinvillei n. sp., jaws with denticulate front borders can also be found in thè Triassic species E. orobicus and E. triasicus from thè Italian Prealps and Grès à Voltzia respectively, as well as in thè Jurassic species E.
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
17
dentatus, thè latter considered by Kozur (1971) as a syno- nym of E. proavus. However, in contrast to E. joinviUei n. sp., all these Eunicites possess smooth surfaced man- dibular processes (Ehlers, 1869; Gali & Grauvogel, 1966; Alessandrello & Bracchi, 2004). Moreover, even if thè subelliptical jaws of E. joinviUei n. sp., E. orobicus and E. triasicus are quite similar in shape, thè jaws of E. joinviUei n. sp. bear 4 teeth, whereas thè same structures in E. oro¬ bicus are equipped with 5-6 teeth (Alessandrello & Brac¬ chi, 2004) and those of E. triasicus with 8-9 teeth (Gali & Grauvogel, 1966). Finally, thè jaws of E. dentatus are scythe shaped and equipped with 6 teeth (Ehlers, 1868).
Eunicites falcatus n. sp.
PI. II (Figs. A, B)
Diagnosis : scythe shaped jaws positioned side by side along a centrai junction line and protruding beyond thè lateral borders of thè forceps. Jaws inclined at an angle of almost 1 80°, with concave and smooth front borders, markedly sharpened lateral ends, straight and tumed for- ward posterior borders. Short and triangular mandibular processes joined for about 1/5 of their length. Very short triangular and symmetrical carriers.
Derivatio nominis : from thè Latin falcatus (= scythe shaped), referring to thè shape of thè jaws.
Geologica l age : Cenomanian.
Type locality : Haqel.
Material : 1 specimen. MSNM Ì8488 (holotype).
Description : thè examined specimen is whole-preserved (120 mm in length) and it has a Constant body width, apart from a slight broadening at thè level of thè pharyngeal region.
The stomodaeal apparatus is 8 mm in length and is mostly preserved as an imprint.
Only thè left jaw is full-preserved, while thè right one is rather damaged and lacks thè anterior pari. The left jaw is 0.7 mm long and 2 mm wide.
The basai parts of thè mandibular processes are quite mineralized while thè other sectors are preserved as weak imprints.
Elongate, arcuate and hooked forceps are articulated to very short triangular carriers. The articulation hinge between carriers and forceps shows a concave pattern. The hooked shape of thè left forcep may be clearly distin- guished and thè imprint of thè extemal margin of thè right forcep is also evident.
At thè mid-length of thè pharyngeal apparatus thè left maxilla II is preserved as a faint subrectangular trace.
The posterior tip of thè left mandibular process of thè holotype is superimposed on thè left maxilla II: this sug- gests a dorsal preservation as in thè eunicidans thè jaw apparatus occupies a ventral position inside thè pharynx, while thè maxillae are dorsally placed (Fauchald, 1992; Fauchald & Rouse, 1997).
Only a few metameres preserve thè aciculae: they appear grouped in bundles of two equally thicknened ele- ments for each parapodium however, it is impossible to establish for certain if they are notopodial or neuropodial. Although in consideration of thè dorsal preservation of thè specimen, they should be interpreted as notopodial. The poor quality of preservation did not allow thè exact number of metameres to be counted.
Remarks : thè complex architecture of thè pharyngeal armature of thè examined specimen clearly marks it as an eunicidan polychaete. Moreover, thè pattern of thè jaws, thè shape of thè paired carriers and thè kind of preserva¬ tion of thè pharyngeal apparatus, formed of a white and pearly material quite similar to thè originai one, suggest that it is a labidognath eunicid or lumbrinerid (Fauchald, 1992; Fauchald & Rouse, 1997). Even if thè eunicids possess, differently from thè lumbrinerids, prostomial antennae that are not evident in thè material examined (George & Hartman-Schròder, 1985), it is preferred here to assign thè specimen to Eunicites , especially due to thè strong similarity of thè stomodaeal armature (Ehlers, 1868, 1869). The jaws of thè examined material are very similar in shape to those of E. affinis and E. dentatus, especially regarding thè sharpened lateral ends (Ehlers, 1868; Alessandrello, 1990) that produce a scythe shape. However, when compared with E. falcatus n. sp., thè mandibular processes of both E. affinis and E. dentatus are longer and stronger. The mandibular processes of both E. affinis and E. dentatus are joined for almost their whole length: in E. falcatus n. sp. thè same structures are joined for about 1/5 of their length. Moreover, thè anterior bor¬ ders of thè jaws are denticulate in both E. affinis and E. dentatus, not smooth like those of E. falcatus n. sp. Jaws with sharpened lateral ends that produce a scythe-shape structure can also be found in E. atavus and in E. proavus. However, thè anterior borders of thè jaws of E. atavus are not simply concave like those of E. joinviUei n. sp., they are undulate, concave extemally and convex intemally. Finally, thè jaws of E. proavus are quite similar to those of E. joinviUei n. sp. with regard both to thè shape and to thè pattern of thè anterior borders, but thè angle they form has a reduced magnitude and thè mandibular processes are triangular, thin, elongate and completely separated.
Eunicites diopatroides n. sp.
PI. II (Figs. C, D); PI. Ili (Figs. A, B); PI. XV
Diagnosis : subelliptical jaws totally joined along a centrai junction line, inclined at an angle of almost 180° and projecting beyond thè lateral borders of thè mandibu¬ lar processes but not protruding more than thè forceps. Jaws anterior and posterior borders smooth and slightly concave, lateral borders markly rounded. Triangular, elongate and grooved mandibular processes, positioned side by side along a median line for about 2/3 of their length. Triangular, short, broad and symmetric carriers completely joined along a median line. Each carrier later- ally provided with a strong and semicircular paddle.
Derivatio nominis: referring to thè resemblance between thè maxillary apparatus of thè examined material and that of thè modem onuphid genus Diopatra.
Geological age: Cenomanian.
Type locality: Haqel.
Material: 3 specimens. MSNM Ì23084 (holotype), MSNM il 3288 (part and counterpart) and MSNM Ì20613 (paratypes).
Description: each specimen lacks thè terminal part of thè body however, thè pharyngeal apparatus is always preserved. It measures 7.8 mm in MSNM 13288, 7 mm in MSNM 20613 and 9 mm in MSNM Ì23084. In MSNM 23084 (132 mm in length) jaws, forceps, fragments of
18
GIACOMO BRACCHI & ANNA ALESSANDRELLO
mandibular processes and carriers are clearly mineralized, while in MSNM 20613 (140 mm in length) only thè pad- dles of thè carriers and thè basai parts of thè mandibular processes are three-dimensionally preserved. Regard- ing MSNM il 3288 (18 mm in length) both thè part and counterpart of thè specimen are available: thè jaws are not preserved in thè part, while thè other components of thè stomodaeal apparatus appear disjuncted and fragmented both in thè part and counterpart. The body shows a Con¬ stant width in MSNM i 1 3288 (8.7 mm) and MSNM 23084 (2.8 mm), while MSNM 20613 (2.8 mm) shows a slight widening at thè level of thè pharyngeal region (3.6 mm).
In MSNM il 3288 thè jaws are slightly distorted, in MSNM 20613 they are also disarticulated. The grooves of thè mandibular processes are evident in thè holotype, thè only specimen showing full-preserved forceps. The for- ceps of thè holotype are thick, notably projecting beyond thè lateral borders of thè jaws and measure 5 mm in length: thè maximum distance between thè extemal margins is 3.7 mm. In MSNM Ì20613 thè imprints of thè forceps are 4.4 mm apart in thè point of maximum separation.
The maxillary apparatus is also represented by well- preserved carriers, a small fragment of thè right maxilla II (MNSNM Ì23084) and a weak imprint of thè left maxilla II (MSNM il 3288). Both thè carriers are laterally pro- vided with a rounded and semicircular device (paddle).
In MSNM il 3288 each parapodium is provided with two strong aciculae tumed toward thè cephalic extrem- ity and one shorter intercalary acicula, divaricating from thè others but similarly oriented. In MSNM Ì23084 thè first half of thè body bears tufts of two equally developed aciculae, while thè other tufts consist of three elements arranged as in MSNM il 3288. The longest aciculae meas¬ ure between 2 mm (MSNM Ì23084) and 3 mm (MSNM il 3288). In MSNM Ì20613 thè aciculae are not preserved. Therefore, as in thè living genus Eunice, thè slender acicula could be interpreted as notopodial, thè thick ones as neuropodial (Fauchald, 1992). The interpretation of thè parapodial position of thè aciculae of MSNM il 3288 and MSNM Ì23084 is confirmed by thè position in which these specimens are preserved: considering thè superim- position of thè jaw apparatus upon thè maxillary appara¬ tus thè ventral surface is evidently being observed.
Some metameres are evident in thè holotype (140: each 1.7 mm in thickness) and MSNM il 3288 (27: each 0.4 mm in thickness).
Apart from short lengths of thè alimentary canal and body outline, thè soft parts are not preserved.
Remarks : thè complex architecture of thè pharyngeal armature of thè examined specimens clearly indicates that they are eunicidan polychaetes. Moreover, thè pat¬ tern of thè jaws, thè shape and architecture of thè paired carriers and thè kind of preservation of thè pharyngeal apparatus, sometimes formed of a white and pearly mate¬ rial quite similar to thè originai one, suggest that they are labidognath eunicids or lumbrinerids (Fauchald, 1992; Fauchald & Rouse, 1997). Even if thè eunicids possess, in contrast to thè lumbrinerids, prostomial antennae that are not evident in thè material examined (George & Hart- man-Schròder, 1985), it is preferred here to assign thè latter to Eunicites, especially due to thè strong similarity of thè stomodaeal armature (Ehlers, 1868, 1869). Subel- liptical jaws and grooved mandibular processes such as those of E. diopatroides n. sp. are typical of E. proavus,
E. joinvillei n. sp. and E. ciffinis. However, thè jaws of both thè latter taxa do not have smooth borders but have a denticulate anterior margin. Moreover, all these species lack carriers laterally equipped with semicircular paddles like those of E. diopatroides n. sp. (Ehlers, 1868; Kozur, 1970; Alessandrello, 1990). Such carriers have already been observed in both modem and fossi 1 eunicemorph polychaetes, such as thè Carboniferous eunicid Esconites zeìus, whose jaws have a denticulate anterior border (Thompson & Johnson, 1977), and thè Jurassic species Melanoraphia maculata. The latter was assigned to thè lumbrinerids even if its affinities within thè eunicidans must stili be demonstrated (Arduini et al., 1982).
Among thè modem eunicids carriers equipped with such paddles can be found, for instance, in some spe¬ cies of Eunice and Marphysa Quatrefages, 1865 (Fauvel, 1923; Lange, 1950; Fauchald, 1970, 1992). Moreover, a pharyngeal apparatus very similar in thè shape and arrangement of thè jaws, mandibular processes and car¬ riers to that seen in E. diopatroides n. sp. can also be found in some species of thè onuphid genus Diopatra Audouin & Milne-Edwards, 1833 (Flartman, 1944; Kozlowski, 1956; Fauchald, 1977). Even if thè onuphids are phylogenetically and morphologically related to thè eunicids, they are tubicolous polychaetes (Paxton, 1986; Orensanz, 1990) that should have left traces of their tubes in thè Haqel limestones. Since tubicolous polychaetes or their tubes have not been found to date in thè fossilifer- ous outcrops of Lebanon, it is preferred here to assign thè examined specimens to Eunicites.
Eunicites mariacristinae n. sp.
PI. Ili (Figs. C, D)
Diagnosis : smooth bordered and half-ellipse shaped jaws, closely positioned side by side along a centrai junction line and forming a V-shaped complex. Elongate, triangular, thin and completely disjuncted mandibular processes.
Derivatio nominis : dedicated to Maria Cristina Gambi, an Italian marine biologist, in recognition of her studies on thè modem polychaetes.
Geologica! age : Cenomanian.
Type locality : Haqel.
Material : 1 specimen. MSNM Ì8490 (holotype).
Description: thè body of thè examined specimen is preserved as a dark, coiled and 110 mm long trace. No mor- phological features were observed because of thè poor qual- ity of preservation. The body has a Constant width of about 2.5 mm, with thè exception of thè pharyngeal region which broadens slightly (3.5 mm). The chaetae are not preserved.
With regard to thè stomodaeal apparatus, only thè jaws and mandibular processes are preserved: they are evident as imprints at a distance of about 1 0 mm from thè base where thè pharyngeal region widens. The jaws show straight ante¬ rior and posterior borders, while thè lateral ones are notice- ably rounded. The lateral borders are quite tumed anteriorly so that thè jaws are inclined at an angle of about 60-70° and they form a V-shaped complex. The internai border of each jaw is straight and 0.15 mm long: it completely coincides j with thè internai border of thè other jaw.
Due to thè absence of both thè chaetae and maxillary apparatus it is impossible to recognize thè position in which thè specimen is preserved.
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA. POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
19
Remarks : thè architecture of thè pharyngeal armature of thè examined specimen clearly indicates that it is an euni- cidan polychaete. Even if thè eunicids possess, in contrast to thè lumbrinerids, prostomial antennae that are not evident in thè material examined (George & Hartman-Schròder, 1985), it is preferred here to assign thè latter to Eunicites, especially due to thè strong similarity in thè pattern of thè jaws (Ehlers, 1 868, 1 869). Among thè species assigned to thè genus Eunicites to date, three show some similarities with E. mariacristinae n. sp. with regard to thè architecture of thè jaws. They are thè Eocenic species E. gazolae and E. pinnai fforn Monte Bolca (Alessandrello, 1990b) together with thè Cretaceous species E. phoenicius ffom Haqel (Alessandrello & Teruzzi, 1986). However, in all these species thè complex formed by thè jaws does not show thè sanie combination of characters as described for E. mariac¬ ristinae n. sp. In E. gazoale and E. phoenicius thè jaws are not half-ellipse shaped and closely positioned side by side, they are instead triangular and joined for a short distance along a centrai junction line. Moreover, in E. gazolae thè lateral extremities of thè jaws are tumed laterally, so that thè jaws form an angle of greater magnitude when compared with that formed by thè same structures in E. mariacristinae n. sp. In conclusion, even if thè mandibular processes of E. pinnai are quite similar to those of E. mariacristinae n. sp., thè jaws of thè former are subtriangular, laterally elongate, slightly arcuate at thè posterior end and, as in E. phoenicius, they form an angle of almost 180°.
Family Lumbrineridae Savigny, 1818 Genus Teruzzia n. gen.
Diagnosis : thread-like and small-bodied worms. Body sometimes tapering both anteriorly and posteriorly. Three pattems recognized in thè morphology of thè jaw apparatus: small, ovai and separated jaws provided with long and triangular mandibular processes joined for about 3/4 of their length; subelliptical jaws fused in a V-shaped complex equipped with long and triangular mandibular processes joined for about 3/4 of their length; jaws and mandibular processes fused in a Y-shaped complex. Max- illae consisting of 5 paired plates as maximum. Short and symmetrical carriers heavily linked to strong and arcuate forceps and sometimes provided with a support, which articulates them to thè maxillae I.
Derivatio nominis : dedicated to Giorgio Teruzzi, for his useful advice and support during this work.
Type species : Teruzzia pezzo/ii n. sp.
Teruzzia pezzo lii n. sp.
PI. IV; PI. XVI
Diagnosis : small and ovai jaws, smooth bordered, concentrically grooved on their surfaces, not joined along a median line. Sub-triangular mandibular processes, longitudinally grooved and joined for about 4/5 of their length but rather divaricate at thè posterior end. Semi- circular carriers equipped with a rounded support, which articulates them to thè forceps.
Derivatio nominis : dedicated to Luigi Pezzoli for his support during this study.
Geologica l age\ Cenomanian.
Type locality : Haqel.
Material. 52 specimens. MSNM il 3290 (holotype), MSNM Ì20611 and MSNM Ì24943 (paratypes), MSNM Ì8342, MSNM Ì8344, MSNM Ì8346, MSNM Ì8382, MSNM Ì8384, MSNM Ì8385, MSNM 8479, MSNM Ì8487, MSNM i9 1 78, MSNM Ì9184, MSNM Ì9193, MSNM Ì9362, MSNM il 2401, MSNM il 2407, MSNM il 2408 (part) and MSNM il 24 12 (counterpart), MSNM il 24 15 (part) and MSNM il 3305 (counterpart), MSNM il 24 19, MSNM il 3284, MSNM il 3297, MSNM il 3298, MSNM il 3299, MSNM Ì13301, MSNM Ì13302, MSNM Ì16282, MSNM Ì20600, MSNM Ì20608, MSNM Ì20628, MSNM Ì20640, MSNM Ì20651, MSNM Ì20653, MSNM Ì20654, MSNM Ì20659, MSNM Ì22836, MSNM Ì23046, MSNM Ì23417 I, MSNM Ì23417 II, MSNM Ì23419 I, MSNM Ì23419 II, MSNM Ì23457, MSNM Ì24926 (part and coun¬ terpart), MSNM Ì24930, MSNM Ì24933, MSNM Ì24936 I (part and counterpart), MSNM Ì24977, MSNM Ì25000, MSNM Ì25001, MSNM Ì25004 and MSNM Ì25438.
Description: thread-like worms, measuring from 20 mm (MSNM Ì23419 I) to 85 mm (MSNM il 2408 and MSNM il 24 12) in length, from 0.2 mm (MSNM il 3284, MSNM Ì23419 I, MSNM Ì23419 II and MSNM Ì24977) to 1.2 mm (MSNM il 3299) in width. The metameres are between 0.1 mm (MSNM Ì20651) and 0.5 mm (MSNM il 2408 and il 24 12) long. In most of thè examined speci¬ mens thè pharyngeal region is not preserved, probably due to thè cuticle slendemess in this body sector.
The complex formed of thè jaws and their proc¬ esses, which are often thè only preserved elements of thè stomodaeal apparatus. It is between 0.8 mm (MSNM Ì8342, MSNM il 3290 and MSNM il 3299) and 1.5 mm (MSNM il 2407) in length. In most of thè examined samples thè jaws are three-dimensionally preserved: they show smooth anterior borders and they protrude slightly beyond thè lateral borders of thè mandibular processes but they are less jutted than thè forceps.
The mandibular processes are longitudinally grooved, elongate, thin and quite expanded behind thè jaws.
The forceps are rather shorter than thè mandibular processes: they measure from 0.4 mm (MSNM Ì25004) to 0.7 mm (MSNM Ì2061 1 and MSNM Ì23457) in length and their apical hooks extend to about 3/4 of thè length of thè mandibular processes. The extemal margins of thè forceps are between 0.2 mm (MSNM Ì25004) and 0.6 mm (MSNM Ì23419 I) apart.
Some specimens (MSNM Ì8342, MSNM Ì8384, MSNM Ì8479, MSNM il 2401, MSNM il 2407, MSNM i 1 3284, MSNM Ì13290, MSNM Ì13298, MSNM Ì13301, MSNM Ì20611, MSNM Ì20628, MSNM Ì20640, MSNM Ì23419 II, MSNM 24943, MSNM 25000, MSNM Ì25001 and MSNM Ì25438) preserve fragments of thè maxil¬ lae II. These structures show a denticulate inner margin provided with 4 teeth. Only MSNM Ì20640 preserves fragments of a maxillary piate which may be a maxilla III, IV or V.
Short (about 0.3 mm), semicircular and symmetrical carriers completely joined along a median line are pre¬ served in most of thè examined specimens. Each carrier is linked on a support, which articulates it to thè forcep. These supports are joined along a median line and have thè shape of a circumference quarter, with a rounded external border and a straight inner margin. In some specimens
20
GIACOMO BRACCHI & ANNA ALESSANDRELLO
(MSNM Ì8479, MSNM il 3290 and MSNM Ì25001 for example) thè carriers appear heavily linked to thè forceps.
Only a few aciculae were observed but they are poorly preserved and it was not therefore possible to investigate how they relate to each parapodium. The aciculae meas- ure from 0.3 mm (MSNM Ì20653) to 0.7 mm (MSNM Ì8385) in length.
With regard to thè soft parts, thè body outline is often preserved. Moreover, thè dark traces of thè alimentary canal are evident in a few specimens. MSNM il 3301 has two anal cirri preserved as thin and faint imprints that measure 1 mm in length.
The dorsal surface is exposed in most of thè specimens examined (for example MSNM Ì8479, MSNM il 3290, MSNM Ì24943 and MSNM Ì25001): this can be clearly deduced from thè position in which thè jaw and maxillary apparata are preserved (Fauchald & Rouse, 1997).
Remarks'. thè generai body morphology together with thè architecture of thè jaw apparatus clearly indicate that Teruzzia n. gen. belongs to thè eunicidans. Furthennore, thè presence of carriers which are heavily linked to thè forceps suggest a labidognath architecture for thè pha- ryngeal apparatus of Teruzzia n. gen. The labidognath eunicidans include Onuphidae, Dorvilleidae, Eunicidae and Lumbrineridae. The assignment of Teruzzia n. gen. to Onuphidae and Dorvilleidae is excluded here. In thè first place, because thè onuphids are tubicolous polychaetes, not free-living forms like Teruzzia n. gen., in thè second because thè maxillary apparatus of thè dorvilleids consists of 2-4 longitudinal rows of small toothed plates and not of 5 paired plates as maximum such as is present in thè Euni¬ cidae, Lumbrineridae and, as it seems, also in Teruzzia n. gen. The distinction between thè lumbrinerids and eunicids is especially based on thè presence of prostomial antennae, non-mineralized structures which are difficult to observe in fossils. However, thè jaw apparatus of Ter¬ uzzia n. gen. is quite similar to that of some species of thè living lumbrinerid Lumbrineris Blainville, 1828 and that of thè Jurassic Lumbriconereites (Ehlers, 1868; Hartman, 1944; Imajima & Higuchi, 1975; Orensanz, 1990). Differ- ently from Teruzzia n. gen., thè jaws of Lumbriconereites (430 mm) are subrectangular and have slightly rounded anterior borders. They are joined along a centrai junction line so that they form an heart-shaped complex.
When Teruzzia n. gen is compared with thè Jurassic species Melanoraphia many differences may be noted. In contrast to thè small and ovai jaws of Teruzzia n. gen, thè jaws of Melanoraphia are triangular and wing shaped. Moreover, thè jaws of Melanoraphia bear short, subtri- angular, rounded and separated mandibular processes (Arduini et al., 1982), rather different from thè complex thè same structures form in Teruzzia n. gen. Finally, thè carriers of Melanoraphia are laterally provided with two semicir- cular paddles that are not present in Teruzzia n. gen.
There are also significant differences between thè lum¬ brinerid Phiops of Mississipian age and Teruzzia n. gen., especially with regard to thè shape of thè jaws (Schram, 1979). The jaws of Phiops are not ovai like those of Ter¬ uzzia n. gen., but are subtriangular, laterally expanded and completely joined along a centrai junction line, with rather concave lateral borders, convex front borders and quite sharpened lateral ends. Moreover, thè carriers of Phiops aciculorum are subrectangular and not semiciur- cular like those of T. pezzolii n. gen. n. sp.
T. pezzolii n. gen. n. sp. and T. sagittifera n. gen. n. sp. may be distinguished on thè basis of thè shape and devel- opment of thè carriers: in thè former they are semicircular and articulated to thè forceps by a rounded support, in thè latter they form an arrow head shaped complex and lack any kind of support. The distinction between T. pezzolii n. gen. n. sp. and T. pusilla n. gen. n. sp. can be limited to thè shape of thè jaws: ovai and separated in thè former, subelliptical and fused in a V-shaped complex in thè latter. Finally, T. pezzolii n. gen. n. sp. and T. gryphoeides n. gen. n. sp. may also be easily distinguished on thè basis of thè morphology of thè jaw apparatus. This consists of two strong subtrapezoidal jaws forming a Y-shaped complex in thè latter and two small ovai jaws hearing triangular, elongate and joined mandibular processes in thè former.
Teruzzia sagittifera n. sp.
PI. V
Diagnosis : small, ovai and smooth-bordered jaws, completely separated and laterally elongate beyond thè extemal borders of thè forceps. Slightly elongate carriers, joined for 1/2 of their length and forming an arrow head shaped complex.
Derivatio nominis : from thè Latin sagittiferus (= hearing an arrow), referring to thè shape of thè com¬ plex formed by thè maxillary carriers.
Geological age: Cenomanian.
Type locality : Flaqel.
Material : 2 specimens. MSNM Ì23436 (holotype) and MSNM Ì8476 (paratype).
Description: thè holotype is flattened on thè surface of a small and thin limestone slab where two fish are also preserved. In both thè holotype and paratype thè body is completely preserved as a rust coloured trace and it is quite coiled: however, a length of 50 mm and a width of 0.5 mm is reasonably valuable. The metameres and acicu¬ lae are not preserved.
The stomodaeal apparatus is 1.3 mm in length and is ; equipped with two elongate, triangular and symmetrical carriers (0.6 mm). Only thè jaws and carriers are com¬ pletely preserved, thè former three-dimensionally, thè latter as imprints. The basai parts of thè forceps are strongly fragmented. Only thè posterior extremities of thè smooth surfaced mandibular processes are preserved as imprints: even if thè mandibular processes are ffagmentary, thè pat¬ tern seems to be very similar to that of thè other species described here under Teruzzia n. gen. In thè holotype, thè superimposition of thè maxillary apparatus on thè mandibu¬ lar processes allow its dorsal preservation to be recognized.
The carriers are heavily linked to two strong and 0.25 mm thick forceps: thè anterior extremities of thè forceps are preserved as arcuate and hooked imprints. The tips of thè forceps do not extend as far as thè posterior borders of thè jaws.
In thè paratype only thè jaws, mandibular processes, carriers are preserved as well as a long reddish trace of , thè body at thè posterior end of thè pharyngeal region: thè superimposition of thè forceps on thè mandibular proc¬ esses allow to recognize thè ventral preservation of this specimen.
Remarks : thè shape of thè jaws and their processes I suggest that thè examined specimen should be assigned to
PALEODrVERSITY OF THE FREE-LIVTNG POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
Teruzzia n. gen. Even if thè jaw apparatus of T. sagittifera n. gen. n. sp. is very similar to that of T. pezzolii n. gen. n. sp., they may be distinguished on thè basis of thè shape of thè carriers. In T. sagittifera n. gen. n. sp. thè carriers are arrow head shaped and are directly articulated to thè forceps, while in T. pezzolii n. gen. n. sp. there are two semicircular supports that articulate thè triangular carri¬ ers to thè forceps. In thè only specimen examined of T. gryphoeides n. gen n. sp. thè carriers are not preserved but thè large, subtrapezoidal jaws are very different from thè small ovai ones of T. sagittifera n. gen. n. sp. The distinc- tion between T. sagittifera n. gen. n. sp. and T. pusilla n. gen. n. sp. can be limited to thè shape of thè jaws: ovai and disjuncted in thè former, subellitptical and fused in a V-shaped complex in thè latter.
Teruzzia pusilla n. sp.
PI. VI (Figs. A, B)
Diagnosis : mandibular processes thick, triangular, elongate, longitudinally grooved, joined for about 4/5 of their length and slightly divaricate posteriorly. Com- pletely fused, slightly elongate and subelliptical jaws forming a V-shaped and smooth bordered complex. Jaws embedded with grooves running parallel to their lateral and posterior borders.
Derivatio nominis : from thè Latin pusillus (= tiny), referring to thè dimensions of thè examined specimen in comparison with thè other species of thè same genus.
Geological age\ Cenomanian.
Type locality : Haqel.
Material : 1 specimen. MSNM Ì8379 (holotype).
Description : thread-like worm that preserves thè cephalic extremity and thè pharyngeal apparatus but lacks thè caudal extremity. At thè level of thè pharyngeal region a length of approximately 3.5 mm of thè body is not preserved. The examined specimen measures 2.5 mm in length, 0.6 mm in width. The latter is Constant. Both sides of thè body are provided with aciculae however, due to thè poor quality of preservation it was not possible to determine thè relationship to each parapodium.
The pharyngeal apparatus appears formed of a pair of joined jaws (0.4 mm in length and 0.25 mm in width) with a smooth anterior border. The junction line between thè jaws is not present so that they seem to form a single V-shaped complex. Two thick mandibular processes are articulated on thè posterior end of thè jaws: they are 0.7 mm in length.
The other elements that comprise thè buccal apparatus are not preserved: thus it was not possible to recognize with certainty thè position in which thè specimen exam¬ ined is preserved.
Discussion : thè mandibular processes of thè examined specimen are very similar in shape and development to those of T. pezzolii n. gen. n. sp. and T. sagittifera n. gen. n. sp. Moreover, thè mandibular processes of T. pusilla n. gen. n. sp. are grooved like those T. pezzolii n. gen. n. sp., even if in T. pusilla n. gen. n. sp. these structures seem more prominent. However, thè subelliptical jaws of thè examined material show a difference in shape when compared with those of T. pezzolii n. gen. n. sp and T. sagittifera n. gen. n. sp.. These possess small, ovai and completely separated jaws which do not form a V-shaped
complex like that of T. pusilla n. gen. n. sp. Finally, in T. gryphoeides n. gen. n. sp. thè jaw apparatus consists only of two subtrapezoidal and elongate jaws that form a Y- shaped complex. Furthermore, thè species included in thè living Lumbrineris can also have, in thè same way as thè strictly related Teruzzia n. gen., a jaw apparatus either with thè pattern of T. pezzolii n. gen. n. sp and T. sagittifera n. gen. n. sp., of T. pusilla n. gen. n. sp. or of T. gryphoeides n. gen. n. sp. (George & Hartman-Schròder, 1985).
Teruzzia gryphoeides n. sp.
PI. VI (Figs. C, D)
Diagnosis : subtrapezoidal and smooth surfaced jaws with denticulate anterior borders, joined along a median line and quite divaricate both anteriorly and posteriorly.
Derivatio nominis : from thè Greek ypi(po£i5e<; (= enig- matical), referring to thè vicissitudes during thè systematic defìnition of thè examined specimens (see thè remarks).
Geological period : Cenomanian.
Type locality. Haqel.
Material : 1 specimen. MSNM Ì24967 (holotype).
Description : thè studied specimen is 35 mm in length and thè body is thread-like. The jaw apparatus and thè caudal extremity are preserved but not thè maxillary apparatus, except for a short length of thè left maxillae I and II. At thè level of thè pharyngeal region thè body is not preserved for a length of about 3.5 mm. The body measures 0.7 mm in width at thè mid-length, but this decreases toward both thè cephalic and caudal extremi- ties. Alongside thè body some imprints of poorly pre¬ served aciculae are evident. The observed elements of thè stomodaeal apparatus are preserved in three-dimensions with thè exception of thè left forcep, which is seen as an imprint.
The jaws measure 1.5 mm in length and are subtrap¬ ezoidal in shape. These subtrapezoidal jaws are joined along their straight and shorter flanks and have concave lateral borders. The anterior and posterior extremities of thè jaws are sharpened and quite divaricate so that thè jaw apparatus appears Y-shaped. On thè side of thè left jaw a 0.5 mm a long imprint of a hooked forcep is preserved together with a longitudinally elongate and subrectangu- lar piate that probably represents a fragment of thè left maxilla II. The latter is partially superimposed on thè underlying jaws that in thè eunicidan polychaetes are typically ventral: a dorsal preservation of thè examined specimen may thus be inferred.
The carriers and chaetae are not preserved.
Remarks : thè morphological pattern of thè jaw appara¬ tus of T. gryphoeides n. gen. n. sp. is very different from all thè other species described here under Teruzzia n. gen. The jaw apparatus of T. gryphoeides n. gen. n. sp. actually lacks thè distinction between jaws and mandibular proc¬ esses that can be so clearly observed in thè other species of thè genus: these structures seem completely fused in a single Y-shaped complex so that thè latter appears formed of two subtrapezoidal jaws only. The pharyngeal armature of thè examined specimen is worthy of note due also to thè absence of carriers.
Among thè eunicemorph polychaetes, a jaw appara¬ tus like that described for T. gryphoeides n. gen. n. sp., or
22
GIACOMO BRACCHI & ANNA ALESSANDRELLO
rather a jaw apparatus formed of two large ventral plates that form a Y-shaped complex, can also be found in thè oenonids and dorvilleids. Moreover, several dorvil- leids possess a ctenognath maxillary apparatus, among thè modera eunicidans this is thè only one that can be characterized by thè absence of carriers, a structure not present in thè examined specimen. However, thè assig- ment of thè examined material to these families may reasonably be excluded. In thè oenonids thè pharynx is armoured with a prionognath maxillary apparatus which typically has paired, slender and elongate carriers and is formed of a heavily sclerotinized, shiny and black mate¬ rial strenghtened with metals. In thè examined material thè carriers are not present but thè jaw apparatus consists of a very different lighter material. This material is whit- ish in colour, pearly, calcifìed and very similar to that of thè labidognath eunicidans, such as thè lumbrinerids. It also resembles that of thè other specimens assigned here to Teruzzia n. gen. Moreover, both in thè prionognath oenonids and dorvilleids thè maxillary apparatus consists of longitudinal rows (2-4) of small, subrectangular and denticulate maxillae (Fauchald, 1970, 1977; Fauchald & Rouse, 1997). T. gryphoeides n. gen. n. sp. possesses large, ventral and Y-shaped jaws however, it lacks car¬ riers like those observed in some dorvilleids. A longitu- dinally elongate and subrectangular fragment of thè left maxilla II is also preserved. It may be inferred from a study of thè morphology of this fragment that thè maxil¬ lary apparatus of T. gryphoeides n. gen. n. sp. shows a very different pattern when compared with those typical of both thè oenonids and dorvilleids. On thè contrary, thè pattern of this fragment seems to fit better with thè maxillae typical of thè labidognath eunicidans, among which only thè lumbrinerids possess a jaw apparatus that consists of a large, ventral and Y-shaped complex like that of T. grypheoides n. gen. n. sp. With regard to thè examined material it is preferred here, for thè reasons outlined, to consider thè absence of thè carriers an artifact of fossilization: thus thè studied specimen is tentatively assigned to thè lumbrinerids and to thè genus Teruzzia n. gen. Moreover, thè generai morphology of thè body and thè kind of preservation that characterize thè specimen assigned to T. gryphoeides n. gen. n. sp. are thè same as for thè specimens assigned to thè other spe- cies of thè genus. Furthermore, also thè species included in thè living Lumbrineris may show, in thè same way as thè closely related Teruzzia n. gen., a jaw apparatus either with thè pattern of T. pezzolii n. gen. n. sp and T. sagittifera n. gen. n. sp., thè pattern of T. pusilla n. gen. n. sp. or with thè pattern of T. gryphoeides n. gen. n. sp. (George & Hartman-Schròder, 1985).
The distinction between T. gryphoeides n. gen. n. sp. and thè other species of Teruzzia n. gen. may be limited to thè jaw apparatus, which in T. gryphoeides n. gen. n. sp. is only formed of two large subtrapezoidal pieces fused in a Y-shaped complex. In T. pezzolii n. gen. n. sp. and T. sag¬ ittifera n. gen. n. sp. thè jaws are small, ovai and clearly distinct from thè triangular and elongate mandibular processes that they carry posteriorly. Finally, in T. pusilla n. gen. n. sp. thè jaws are fused in a V-shaped complex but are nevertheless, clearly distinct from thè mandibular processes. The latter are quite similar in shape and devel- opment to those of T. pezzolii n. gen. n. sp. and T. sagit¬ tifera n. gen. n. sp.
Genus Lumbriconereites Ehlers, 1 869
Lumbriconereites hadjulae n. sp.
PI. VII, PI. Vili
Diagnosis'. subrectangular and smooth bordered jaws, spatulate, grooved, anteriorly elongate, positioned side by side for about 3/4 of their length and slightly divaricate anteriorly. Short, slender, triangular and medially joined mandibular processes. Subtriangular and symmetrical carriers forming a arrow head shaped complex, medially joined for approximately 3/4 of their length and poste¬ riorly divaricate at thè back. Presence of a semicircular support, which links thè carriers to thè forceps.
Derivatio nominis : referring to Hadjula, site of finding of two specimens assigned to this new species.
Geologica! period : Cenomanian.
Type locality : Haqel.
Material : 16 specimens. MSNM il 2424 (holotype, part and counterpart), MSNM il 0402, MSNM 12632 (part and counterpart) and MSNM Ì20614 (paratypes), MSNM Ì8489, MSNM 9359 (part and counterpart), MSNM Ì20623, MSNM Ì22831, MSNM Ì23441, MSNM Ì25091, MSNM Ì25092, MSNM i251 15 (part and coun¬ terpart), MSNM Ì25120 (Hadjula), MSNM Ì25125 (Had¬ jula), MSNM Ì25445 and MSNM Ì25446.
Description : only a few specimens (MSNM Ì9359, MSNM Ì20614, MSNM Ì25092, MSNM i251 15, MSNM Ì25120, MSNM Ì25445) are ftdl-preserved. The body measures from 40 mm (MSNM Ì9359) to 240 mm (MSNM Ì25120) in length, from 0.8 mm (MSNM Ì25445) to 4.5 mm (MSNM Ì252120) in width. The segmentation of thè body is clearly evident only in MSNM Ì25091 (94 metameres: 0.3 mm in thickness), MSNM Ì252120 (150 metameres: 0.7 mm in thickness) and MSNM Ì25446 (118 metameres: 0.5 mm in thickness), but thè full number of thè metameres is never preserved.
The pharyngeal armature measures from 4 mm (MSNM Ì8489) to 4.5 mm (MSNM il 2424) in length and has very elongate jaws. The jaws measure from 0.8 mm (MSNM Ì254746) to 1 .8 mm (MSNM Ì9539, MSNM il 0402 and MSNM Ì25120) in length and they protrude slightly beyond thè lateral borders of thè forceps. The lateral borders of thè jaws are concave, whereas both thè posterior and thè anterior borders are convex. The surfaces of thè jaws are characterized by straight growth lines running parallel to thè lateral and posterior borders.
The small mandibular processes are preserved only in two specimens (MSNM il 2632 and MSNM i206 14), where they appear to be shorter than thè jaws, they show slightly rounded borders and are divaricate at thè posterior extremity.
In most of thè examined specimens thè forceps are also evident but their anterior extremity is never pre¬ served. Nevertheless, thè arcuate and hooked shape of thè forceps may often be desumed by thè investigation of their fragments or partial imprints. The presence of two elongate (almost from 1 .5 mm in MSNM Ì8489 to 2.7 mm in MSNM i25 1 20) and slightly distant (0.6 mm in MSNM Ì25445 and MSNM Ì25446, 1.1 mm in MSNM il 2424) structures is observed. The anterior ends of thè forceps seem to extend as far as thè posterior borders of thè jaws.
In most of thè examined specimens some disjuncted remains of thè maxillary apparatus are preserved. For example, between thè forceps of MSNM il 0402, a small
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRJPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
fragment of thè maxilla II is preserved: it is armoured with four small teeth on thè inner border. A small piate with three longer teeth which may be attributed to thè maxilla III is preserved in thè same specimen. A similar situation characterizes thè maxillary apparatus of MSNM Ì9539. Some fragments of thè maxillary apparatus are evi- dent also in thè counterpart of MSNM i 1 2424, in both part and counterpart of MSNM il 2632 as well as in MSNM Ì22831 and MSNM 25092.
The carriers measure from 1 mm (MSNM Ì25446) to 2.6 mm (MSNM il 0402) in length. They are joined for almost their whole length but are divaricate at thè poste- rior extremity where their tips are 1 mm apart. Each car¬ rier is linked to a support that articulates it to thè forcep. These supports are well preserved only in thè paratype, where they form a semicircular complex slightly narrower than thè carriers.
In many specimens thè aciculae appear as faint, scat- tered imprints or they are not preserved. However, at least two aciculae seem to be present in each parapodium. The parapodia have three aciculae only in MSNM Ì25092. The aciculae are always thè same length and thickness and they measure from 1.1 mm (MSNM Ì25445 and MSNM Ì25446) to 1.8 mm (MSNM i25 1 20) in length. Due to thè absence of morphological differences in thè preserved aciculae of each specimen it was not possible to establish whether these structures were notopodial or neuropodial.
The superimposition of thè maxillary apparatus on thè jaw apparatus suggests a dorsal preservation both for thè holotype and paratype.
The preserved soft parts include thè body outline and dark traces of thè alimentary canal.
Remarks : thè complex architecture of thè pharyngeal armature of thè examined specimens clearly indicates that they are eunicidan polychaetes. Moreover, thè pat¬ tern of thè jaws, thè shape and architecture of thè paired carriers and thè kind of preservation of thè pharyngeal apparatus, sometimes formed of a white and pearly mate¬ rial quite similar to thè originai one, suggest that they are labidognath eunicids or lumbrinerids (Fauchald, 1992; Fauchald & Rouse, 1997). However, thè eunicids possess, in contrast to thè lumbrinerids, prostomial antennae that are not evident in thè material examined (George & Hart- man-Schròder, 1985). Even if thè absence of antennae in thè fossil polychaetes could be an artifact of fossilization, thè similarity between thè pharyngeal armature of thè examined specimens and those of thè species included in thè living genus Lumbrineris is remarkable. Thus, it is preferred here to assign thè specimens examined to Lumbriconereites , especially due to thè strong similar¬ ity of thè stomodaeal armature (Ehlers, 1 869). Only one full-preserved specimen has been assigned to Lumbrico¬ nereites to date. It is thè Jurassic species L. deperditus from Solnhofen (Ehlers, 1869). Although thè shape of thè jaws of L. hadjulae n. sp. is very similar to that of thè jaws of L. deperditus and L. garassinoi n. sp., thè latter described below, these three species can be easily distinguished on thè basis of thè structure of thè carriers. The triangular shape of thè carriers of L. deperditus is not as regular as that of L. hadjulae n. sp. This is due to thè fact that in L. depertidtus thè carriers have rounded lateral borders, while thè anterior and inner margins are straight. Consequently thè complex formed by thè carriers of L. deperditus is elongate, thicker and stronger than that of L.
hadjulae n. sp. However, thè semicircular support, which links thè carriers to thè forceps, is thè more signifìcant feature for distinguishing between thè carriers of L. had¬ julae n. sp. and L. deperditus : it is clearly evident in all thè specimens assigned here to L. hadjulae n. sp. but is absent in both L. deperditus and L. garassinoi n. sp.
Lumbriconereites garassinoi n. sp.
PI. IX (Figs. A, B)
Diagnosis : subrectangular and smooth bordered jaws, spatulate, grooved, anteriorly elongate, positioned side by side for about 3/4 of their length and anteriorly divaricate. Very short, slender, subtriangular and medially joined man- dibular processes. Sub-triangular and short carriers with concave lateral borders and sharpened posterior ends.
Derivatio nominis : dedicated to Alessandro Garassino, in recognition of his studies on fossil invertebrates from Lebanon.
Geologica! age : Cenomanian.
Type locality : Haqel.
Material : 1 specimen (holotype). MSNM Ì9355 (part) and MSNM Ì20618 (counterpart).
Description : thè examined specimen lacks thè poste¬ rior part of thè body. The preserved fragment measures of 1 80 mm in length and 3.7 mm in width. Along thè anterior extremity of thè body some faint traces of thè metameres may be observed. In some cases also thè aciculae are pre¬ served along thè two flanks of thè body, however, due to thè poor quality of preservation of thè specimen it was not possible to evaluate thè number of metameres which form thè trunk or thè number of aciculae related to each parapodium.
The stomodaeal apparatus is approximately 6.2 mm long and thè jaws measure 2 mm in length. Some parallel growth lines are preserved as thin ridges on thè surface of thè jaws: these ridges run from thè anterior to thè poste¬ rior border of each jaw.
Two short, triangular mandibular processes are pre¬ served at thè back of thè jaws. The part also has arcuate forceps preserved: thè left forcep is integrally preserved apart from thè anterior hook, but thè hook and thè basai part of thè right forcep are evident. The extemal margins of thè forceps are 1 .6 mm apart.
The short, triangular and medially joined carriers show a slight broadening at thè articulation hinge with thè forceps.
Fragments of thè dorsal maxillae are present both in thè part and counterpart, however due to thè poor qual¬ ity of preservation it is not possible to determine whether they belong to maxilla II, III or IV.
The relative positions of thè jaw and maxillary appa¬ rata suggest a dorsal preservation for thè holotype.
Remarks : thè complex architecture of thè pharyngeal armature of thè examined specimen clearly indicates that it is an eunicidan polychaete. Moreover, thè pattern of thè jaws, thè shape and architecture of thè paired carriers and thè kind of preservation of thè pharyngeal apparatus, formed of a white and pearly material, suggest that they are labidognath eunicids or lumbrinerids (Fauchald, 1992; Fauchald & Rouse, 1997). However, thè eunicids possess, in contrast to thè lumbrinerids, prostomial antennae that are not evident in thè material examined (George & Hart-
24
GIACOMO BRACCHI & ANNA ALESSANDRELLO
man-Schròder, 1985). Even if thè absence of antennae in thè fossi 1 polychaetes could be an artifact of fossilization, thè similarity between thè pharyngeal armature of thè examined specimens and those of thè species included in thè living genus Lumbrineris is remarkable. Thus, it is preferred here to assign thè studied specimen to Lumbri- conereites, especially due to thè strong similarity of thè stomodaeal armature (Ehlers, 1869). The morphology of thè jaw apparatus of L. garassinoi n. sp. is very similar to that of L. hadjulae n. sp., except in thè shape of thè carri- ers which justifìes thè assignment of thè examined speci¬ men to a different species. Moreover, in contrast to L. hadjulae n. sp., thè maxillary apparatus of L. garassinoi n. sp. lacks, as does that of L. deperditus, any kind of sup- port linking thè carriers to thè forceps. Finally, thè carriers of L. deperditus are subtriangular thè same as those of L. garassinoi n. sp., although in thè latter they are rather less elongate. However, their lateral borders are convex, not concave.
Family Oenonidae Kingberg, 1965 Genus Phoeniciarabella n. gen.
Diagnosis : elongate and thin worms. Jaws from sub¬ triangular to subelliptical, sometimes fiised in a single subtrapezoidal or heart-shaped complex, sometimes sepa- rated. Smooth surfaced or grooved mandibular processes, from subtriangular to subrectangular. Very long, slender and paired carriers, apparently not associated to a median unpaired piece, linked to thè forceps for a short distance, more or less joined along a centrai junction line, often more elongate than thè remaining pharyngeal apparatus and some¬ times laterally provided with a small pinnula. Forceps with denticulate inner borders. Five pair of maxillae at least.
Derivatio nominis : from thè Latin phoenicius (= native of Phoenicia , thè latin name for modem Leba- non) and from Arabella , a living oenonid genus.
Type species : Phoeniciarabella pinnulata n. sp.
Phoeniciarabella pinnulata n. sp.
PI. IX (Figs. C, D)
Diagnosis : subtriangular and elongate jaws, medially joined for about 3/4 of their length, slightly divaricate anteriorly and forming a trapezoidal complex. Triangular, elongate, thick and smooth surfaced mandibular proc¬ esses. Long and slender carriers laterally equipped with a small pinnula and joined along a centrai junction line for about 1/3 of their length.
Derivatio nominis : from thè Latin pinnulatus ( equipped with a paddle), referring to thè presence of a small pinnula on each carrier.
Geologica! age : Cenomanian.
Type locality : Haqel.
Material : 1 specimen. MSNM il 24 11 (holotype).
Decription : thè holotype measures 20 mm in length and 2 mm in width. The aciculae and thè segmentation of thè body are not preserved.
The pharyngeal apparatus is 3.7 mm long and is pre¬ served partly in three-dimensions, partly as an imprint. The maxillary apparatus and thè extemal borders of thè
jaws are mineralized by a black, shiny material that is quite similar to thè originai material and which gives evidence of a heavily sclerotinized pharyngeal apparatus (Fauchald & Rouse, 1997). The jaws are 0.8 mm long and 0.4 mm wide and form a subtrapezoidal complex in which thè shorter side coincides with thè articulation hinge betw een jaws and mandibular processes. The smooth bor¬ ders of thè jaws exhibit a straight pattern apart from thè anterior ends, where a convex pattern is exposed.
Beside thè left jaw, two small, thick, hooked and slightly arcuate structures are evidenti they represent thè left maxilla III and thè left maxilla II respectively. Their tips are oriented backward. Other small plates preserved between thè basai parts of thè forceps can be interpreted as fragments of thè maxillary apparatus.
The right mandibular process is fragmented but clearly evident behind thè jaws. It is short, subtriangular and slightly tapering at thè back extremity. The right man¬ dibular process partially hides thè tip of thè underlying right forcep: this is evidence of ventral preservation. The left forcep is three-dimensionally preserved at thè mid- length only: along its inner margins two small teeth may be distinguished. The forceps are linked to thè carriers for a short distance and are separated with a maximum distance apart of 0.8 mm. Their tips extend as far as thè posterior borders of thè jaws.
The carriers are quite elongate (1.9 mm) and laterally bear an anteriorly directed, small and rounded pinnula, at a point 1/4 of thè distance along their length. These pinnulae have rounded tips and show a slight opening from thè principal axis of thè carriers. The carriers are slightly divaricate at thè posterior extremity and are quite expanded at thè articulation hinge with thè forceps, where they form a semicircular complex with convex lateral bor¬ ders and a concave anterior borders.
The superimposition of thè jaw apparatus on thè max¬ illary apparatus suggests a ventral preservation for thè specimen examined.
Remarks : thè complex jaw apparatus of thè examined specimen clearly indicates that Phoeniciarabella n. gen. is an eunicidan polychaete. The long, slender morphology of thè maxillary carriers, thè forceps linked for a short distance to thè carriers as well as thè heavily sclerotinized jaws and maxillae are all evidence of thè prionognath architecture which characterizes thè pharyngeal appara¬ tus of Phoeniciarabella n. gen. The same characteristics allow Phoeniciarabella n. gen. to be assigned to thè oeno- nids (Colbath, 1986; Fauchald & Rouse, 1997). Among thè modem eunicemorph polychaetes a prionognath sto¬ modaeal apparatus can be found in thè Histriobdellidae Vaillant, 1980 and thè Oenonidae. However, thè former are highly modified, achaetous and tiny polychaetes (< 2 mm) that live as parasites in thè gills of crustaceans. They possess a single unpaired carrier (George & Hart- mann-Schròder, 1985). Moreover, thè carriers of Phoe¬ niciarabella n. gen. are very similar to those of thè living genera Aglaurides Ehlers, 1868, Arabella , Drilonereis, Notocirrus and Oenone (Hartman, 1944; Fauchald, 1970, 1977; George & Hartmann-Schròder, 1985; Orensanz, 1990). The maxillary carriers of these modem genera are usually provided with a third unpaired median piece that is not present in Phoeniciarabella n. gen.: it ts pos- sible that thè absence of this structure in thè examined specimen is a fossilization artifact. However, among thè
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
25
oenonids, this unpaired median piece can be absent in both parasitic (Martin & Britayev, 1998) and free-living taxa (Orensanz, 1990): if thè absence of this structure in Phoeniciarabella n. gen. is reai, this character could sug- gest a primitive condition.
Among thè modem free-living oenonids thè number of maxillae varies within both genera and species (Colbath, 1989) but thè large-bodied forms, such as Arabella and Oenone, usually possess five or more pairs of maxillae, while thè small-bodied forms, such as Drilonereis, have a smaller number of maxillae (Fauchald & Rouse, 1997). The dimensions as well as thè five pairs of maxillae of thè examined specimens suggest that Phoeniciarabella n. gen. is related with thè large-bodied oenonids.
Moreover, P. pinnulata n. gen. n. sp. seems to share some similarity with thè Silurian Synclinophora syncli¬ nalis, recently re-described on thè basis of an articulated apparatus as thè oldest arabellid known (Mierzejewski, 1984): thè arabellids have been recently moved to thè oenonids (Colbath, 1989; Orensanz, 1990). Although S. synclinalis lacks thè pinnulae that provide thè lateral bor- ders of thè carriers of P. pirmulata n. gen. n. sp. and even if it shows maxillae and carriers very similar to those of P. pinnulata n. gen. n. sp., these two species are hardly comparable as in S. synclinalis thè jaws are not preserved. Furthermore, maxillae and carriers have a very similar pattern among thè oenonids so that their generai features are not diagnostic for distinguishing Synclinophora from Phoeniciarabella n. gen.
P. pinnulata n. gen. n. sp. differs from P. caesaris n. gen. n. sp. and P. orensanzi n. gen. n. sp. on thè basis of thè shape of thè jaws and thè pinnulae that occur laterally on each carrier. Neither P. caesaris n. gen. n. sp. nor P orensanzi n. gen. n. sp. possess these pinnulae. Moreover, in contrast to P. pinnulata n. gen. n. sp., thè jaws of P. orensanzi n. gen. n. sp. are subelliptical and separated, are not subtriangular and are fused in a subtrapezoidal com¬ plex, while thè same structures of P caesaris n. gen. n. sp. are subelliptical and form an heart-shaped complex.
Phoeniciarabella orensanzi n. sp.
, PI. X (Figs. A, B)
in width. The jaws are positioned side by side along a mediai line and their rounded lateral ends protrude sig- nificantly beyond thè lateral borders of thè mandibular processes. Behind thè jaws thè 0.6 mm long mandibular processes are preserved. The left jaw and its process are well preserved while thè right ones are quite shattered.
Some fragments of thè mediai and posterior sectors of long and slender carriers are three-dimensionally pre¬ served and appear as shiny, black pieces while thè remain- ing parts are evident as imprints or, as in thè case of thè anterior extremities, are not preserved at all. The carriers are joined along a centrai junction line for about 4/5 of their length and are slightly divaricate at thè posterior extremities. The complex formed by thè carriers is very long ( 1 .5 mm) when compared with thè whole pharyngeal apparatus (about 2.7 mm).
The lack of thè forceps and maxillae did not allow further information to be acquired.
The superimposition of thè mandibular processes on thè maxillary carriers over a short distance suggests a ventral preservation for thè specimen examined.
Remarks : thè assignment of thè specimens examined to Phoeniciarabella n. gen. is mostly justifìed on thè basis of thè shape of thè carriers and thè kind of preservation of thè pharyngeal armature. In thè same way as thè other species of thè genus, thè carriers of P. orensanzi n. gen. n. sp. are long, slender and are not equipped with a third unpaired piece. The pharyngeal armature shows miner- alization by a black, shiny material. Differently from P pinnulata n. gen. n. sp., P. orensanzi n. gen. n. sp. has significanti larger dimensions and differently shaped jaws and carriers. In particular thè jaws of P orensanzi n. gen. n. sp. are subelliptical, not subtriangular, whereas thè carriers do not possess thè pinnulae that characterize those of P pinnulata n. gen. n. sp. As in P. orensanzi n. gen. n. sp., thè carriers in P. caesaris n. gen. n. sp. are also not provided laterally with a pinnula, however, thè jaws of P caesaris n. gen. n. sp. are fused forming an heart- shaped complex. Its mandibular processes are triangular, thick, short and sharpened at thè posterior extremities and longitudinally grooved, in contrast to thè triangular, thinner, longer, smooth-surfaced and posteriorly rounded mandibular processes of P orensanzi n. gen. n. sp.
Diagnosis : stomodaeal apparatus with subelliptical, lat¬ erally elongate and separated jaws. Smooth surfaced, thick and subrectangular mandibular processes with rounded posterior borders. Long and slender carriers joined along a centrai junction line for about 4/5 of their length.
Derivatio nominis : dedicated to José-Maria Orensanz, for his contribution to thè knowledge of oenonid systematics.
Geologica/ age : Cenomanian.
Type locality : Haqel.
Material : 1 specimen. MSNM il 3283 (holotype).
Description : thè preserved body measures 150 mm in length and 5 mm in width at thè mid-length. It lacks thè caudal extremity.
On both sides of thè body a few poorly preserved aciculae are evident as imprints but it is not possible to distinguish their parapodial organization and exact length. The dark trace of thè digestive tube is preserved along thè centrai sector of thè body.
The stomodaeal apparatus is comprised of two smooth bordered jaws that measure 0.2 mm in length and 0.3 mm
Phoeniciarabella caesaris n. sp.
PI. X (Figs. C, D); PI. XI (Figs. A, B)
Diagnosis : stomodaeal apparatus with subelliptical jaws forming an heart-shaped complex. Triangular, thick, short, smooth-surfaced mandibular processes trimed with longitudinal grooves and sharpened at thè posterior extremities. Long and slender carriers, completely joined along a centrai junction line.
Derivatio nominis : in memory of Cesare Alessandrello.
Geologica! age : Cenomanian.
Type locality. Haqel.
Material : 2 specimens. MSNM i206 10 (holotype, part and counterpart) and MSNM Ì23421 (paratype).
Description: instead of thè holotype, it is thè paratype which is completely preserved, however, thè jaw appara¬ tus of MSNM Ì20610 is better preserved. The fragmented body of thè holotype is 270 mm in length and 10 mm in width, while thè filli -preserved paratype measures 60 mm
26
GIACOMO BRACCHI & ANNA ALESSANDRELLO
in length and 0.7 mm in width. The body measurements of both thè holotype and paratype are typical, in thè same way as thè other species of thè genus, of a large-bodied oenonid.
The pharyngeal armature of thè paratype is 1.9 mm long and it is mostly preserved as an imprint.
The jaws are 0.5 mm long and 0.3 mm wide in thè hol¬ otype, 0.25 mm long and 0.15 mm wide in thè paratype. They are smooth-bordered, medially joined along 3/4 of their length, quite divaricate and rounded at thè anterior extremities so that they forni an heart-shaped complex.
Two strong mandibular processes are preserved in both thè considered specimens and they are 1 mm long in thè holotype, 0.35 mm in thè paratype. In thè counterpart of MSNM Ì20610 some longitudinal grooves are preserved on thè surfaces of thè mandibular processes.
In thè paratype, thè extemal margin of thè left forcep and thè basai part of thè right forcep are preserved as imprints. The forceps laterally protrude beyond thè lateral borders of thè jaws and they extend as far as thè posterior borders of thè same structures.
Two very long and slender carriers (3 mm in holotype, 1 . 1 mm in paratype) are linked for a short distance to thè forceps. In thè same way as thè other elements of thè pha¬ ryngeal apparatus, thè carriers are preserved as imprints, apart from a small sector of thè right one and a basai por- tion of both which are formed a black and shiny material.
In thè counterpart of thè holotype some fragments of thè two plates that comprise thè left and thè right maxilla II are preserved.
In thè paratype a few imprints of aciculae appear scattered on both sides of thè body. The poor quality of preservation of thè aciculae did not allow thè number of elements on each metamere to be established.
The superimposition of thè mandibular processes on thè maxillary apparatus suggests a ventral preservation for thè specimens examined.
Remarks : thè assignment of thè specimens examined to Phoeniciarabella n. gen. is mostly justifìed on thè basis of thè shape of thè carriers and thè kind of preservation of thè pharyngeal armature. In thè same way as thè other species of thè genus, thè carriers of P. caesaris n. gen. n. sp. are long, slender and are not equipped with a third unpaired piece. The pharyngeal armature shows miner- alization by a black and shiny material. In P. pinnulata n. gen. n. sp. thè carriers are provided with a lateral pinnula and are joined for about 1/3 of their length, whereas in P. orensanzi n. gen. n. sp. thè carriers lack these pinnulae and are joined for about 4/5 of their length. In P. caesaris n. gen. n. sp. thè carriers do not have any kind of pinnula and are joined along a median line over their entire length. Moreover, P. caesaris n. gen. n. sp. may be distinguished from thè other species of thè same genus by thè shape of thè jaws: subtriangular and fused in a subtrapezoidal com¬ plex with regard to P. pinnulata n. gen. n. sp., subelliptical and separated in P. orensanzi n. gen. n. sp. and subellipti¬ cal and fused in a V-shaped complex in P. caesaris n. gen. n. sp.
Family Didonidae n. fam.
Diagnosis : elongate (50-150 mm) and thick euni- cidans (1.5-3 mm). Jaw apparatus lacking mandibular
processes. White and pearly jaws equipped with a black and shiny maxillary apparatus. Maxillary apparatus consisting of two longitudinal rows of pieces located on both sides of thè pharyngeal armature. Presence of a single, unpaired carrier linked for a short distance to thè forceps.
Remarks : thè assignment of thè specimens described below to a new family within thè eunicemorph polycha- etes is justifìed on thè basis of several characters that determine thè position of this new group as intermediate between thè labidognath oenonoideans such as thè lum- brinerids, thè prionognath eunicidans such as thè histriob- dellids and oenonids, and thè dorvilleids.
The didonids share with thè lumbrinerids thè archi- tecture and conservation of thè jaws. Their jaws have a very similar pattern to that of some living species of thè genus Lumbrineris. They are comprised of a white and pearly material in thè same way as both thè living labidognath eunicidans and those from thè Cenomanian of Lebanon, but differently from thè oenonids. The pha¬ ryngeal armature of thè latter can bear jaws very similar in shape to those of some Lumbrineris and didonids but it is formed of a very different material. However, thè jaw apparatus of thè didonids lacks thè mandibular proc¬ esses that are typically present in thè similarly shaped jaws of thè lumbrinerids, other labidognath eunicidans and thè oenonids. Furthermore, thè carrier of thè dido¬ nids is elongate and has a short linkage with thè forceps, characteristics that are more fitting with thè pharyngeal armature of thè prionognath eunicidans and dorvilleids. Among thè modem prionognath eunicidans only thè histriobdellids possess an unpaired carrier quite similar to that of thè didonids. Neverthless, thè histriobdellids are symbiotic polychaetes that show some adaptations to a parasitic mode of life, such as thè very reduced dimensions, thè absence of chateae and thè presence of two posterior appendixes equipped with a sucker at thè terminal end of thè body. These characteristics together with thè shape of thè jaws indicate that thè histriobdel¬ lids, parasites of cmstacean gills, are very different from thè free-living didonids and from all free-living polychaetes known. The disposition of thè maxillae of thè didonids closely recalls that of both thè prionognath oenonoideans and thè dorvilleids however, their con¬ servation and consistence seem typically labidognath. In thè examined specimens thè maxillary apparatus is preserved by a black, shiny material, very similar to thè heavily sclerotinized material enriched with metal-ions that comprises thè maxillary apparatus of thè prionog¬ nath eunicidans and of thè oenonids from thè Cenoma¬ nian of Lebanon. Furthermore, in thè same way as thè didonids, thè carriers of both thè prionognath eunicidans and dorvilleids show a short linkage with thè forceps. However, thè maxillary apparatus of thè didonids dif- fers from both that of thè labidognath and prionognath eunicidans as it does not include paired and symmetrical carriers but, in thè same way as thè dorvilleids, it bears a single unpaired carrier.
Didone n. gen.
Diagnosis : as thè type species.
Derivatio nominis: dedicated to Didone, mithological
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCR1PTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
27
Queen of thè Phoenicians, ancient inhabitants of Lebanon. Type species : Didone pulcherrima n. sp.
Didone pulcherrima n. sp.
PI. XI (Figs. C, D); PI. XII
Diagnosti: elongate and subelliptical jaws, posteriorly joined for about 3/4 of their length, anteriorly divaricate, forming a V-shaped complex and lacking in mandibular processes. Two longitudinal rows of hooked shaped max- illae. Maxillary apparatus supported by a single elongate and scythe-shaped carrier.
Derivatio nominis : from thè Latin pulcherrimus (= very beautiful), due to thè excellent preservation of thè examined specimens.
Geologica/ age: Cenomanian.
Type locality : Haqel.
Material : 5 specimens. MSNM Ì20617 (holotype, part and counterpart), MSNM Ì20624 and MSNM Ì25457 (paratypes), MSNM Ì20619 and MSNM Ì23415.
Description : apart from MSNM Ì20624, all thè exam¬ ined specimens are preserved for thè entire length of their bodies. They measure from 23 mm (MSNM Ì23415) to 150 mm (MSNM Ì25457) in length. The holotype is 115 mm in length while in MSNM Ì20624 only a 50 mm frag- ment including thè cephalic extremity is preserved. The width of thè body is Constant and it measures from 1.3 mm (MSNM Ì20617) to 35 mm (MSNM i206 1 9), with thè exceptions of MSNM Ì20617 and MSNM Ì25457 that show a significant broadening at thè level of thè pharyn- geal region, from 1.3 mm to 3.5 mm and from 3 mm to 5 mm respectively. The soft parts are not preserved, with thè exception of some short, dark traces of thè alimentary canal and thè body outline.
Each specimen has thè pharyngeal apparatus pre¬ served, thè length reaches 8 mm in MSNM Ì25457, 4.5 mm in MSNM Ì20624, 4 mm in MSNM 20619 and 3.6 mm in MSNM Ì23415. The jaws have smooth borders and longitudinally grooved surfaces. They are 2.2 mm long in MSNM Ì25457, 1.4 mm in MSNM Ì20617, 1.2 mm in MSNM Ì20624 and 1 mm in MSNM Ì20619. The jaw apparata are preserved both as imprints and reliefs.
The maxillary elements are positioned along two lon¬ gitudinal rows on both sides of thè pharyngeal armature. They seem like small hooked-shaped plates whose tips are tumed laterally and slightly backward. In thè holotype at least fìve of these maxillary plates are evident along thè right side of thè stomodaeal apparatus and three along thè left side. Three of these maxillary plates are preserved in thè paratype only along thè right side.
The basai parts of thè forceps are preserved in MSNM Ì20617, MSNM Ì20624 and MSNM Ì25457. In MSNM Ì20624 they appear to be linked for a short distance to thè carrier.
In each specimen thè maxillary apparatus appears to be equipped with a single asymmetrical carrier thè origi¬ nai scythe shape of which may clearly be recognized in thè holotype and MSNM Ì20619.
The aciculae are preserved as imprints along both body flanks of MSNM i203 17, MNSM Ì20624, and MSNM Ì25457. In MSNM Ì25457 they have a length of approximately 2 mm.
As thè jaw apparatus appears superimposed on thè maxillary apparatus in both thè holotype and paratype, thè ventral preservation of these specimens may be implied.
Remarks : thè complex morphology of thè stomodaeal apparatus allows thè examined specimens to be placed within thè Eunicida. The most significant features of thè pharyngeal apparatus are thè absence of thè mandibular processes, thè maxillae arranged in two longitudinal rows and thè presence of an unpaired carrier. Although thè shape of thè jaws of Didone n. gen. resembles that of some lumbrinerids superile ially, thè arrangement of thè maxillae seem typically prionognath. In addition thè carrier is not heavily linked to thè first pair of maxillae as happens in thè click-mechanism of thè lumbrinerid- onuphid-eunicid labidognath line (Fauchald, in lift.) and thè shape of thè carrier is typically dorvilleidean. Thus, Didone n. gen. and thè group to which it belongs may be related to thè oenonid-dorvilleid line.
Order Phyllodocida Dales, 1962 Superfamily Aphroditacea Fauchald, 1977 Family Aphroditidae Malmgren, 1867 Genus Paleoaphrodite Alessandrello & Teruzzi, 1986
Paleoaphrodite libanotica n. sp.
PI. XIII
Diagnosti : small-sized and elliptical body equipped with biramous parapodia hearing long chaetae grouped in two strong tufts. Each tufi: formed of two setal types.
Derivatio nominis : from thè Latin libanoticus (= from Lebanon), referred to country of origin of thè examined material.
Geologica l age : Cenomanian.
Type locality : Haqel.
Material : 6 specimens. MSNM il 6288 (holotype), MSNM Ì23085 and MSNM Ì23086 (paratypes), MSNM i 12397, MSNM Ì23087 and MSNM Ì24937.
Description: thè bodies of thè examined specimens are whole-preserved, apart from MSNM Ì23087 and MSNM Ì24937. The latter two specimens retain only a short frag- ment of thè posterior body part, measuring 23 mm and 8 mm in length respectively. The body measures between 15 mm (MSNM Ì23085) and 20 mm (MSNM Ì23086) in length, between 2 mm (MSNM Ì23085) and 5 mm (MSNM Ì23086) as maximum width. MSNM Ì23086 and MSNM Ì24937 are full-preserved as imprints.
Excluding thè body outline and alimentary canal, thè latter preserved as a dark imprint in MSNM Ì24937, thè soft parts are never evident. Along thè alimentary canal of MSNM Ì24937 some small fìsh vertebrae are preserved: they could be interpreted as traces of predation.
The segmentation of thè body is also not evident even if thè metamerical organization is revealed by thè para- podial arrangement of thè chaetae which have allowed at least 12 metameres in MSNM Ì24937, 16 in MSNM il 6288, 17 in both MSNM il 2397 and MSNM Ì23085 to be counted. The chaetae appear clumped in two tufts on each side of thè metameres: this arrangement is clear evidence of biramous parapodia. Since there are no mor- phological differences among thè chaetae that provide thè tufts of thè same metamere side, it is not possible to dis¬ tinguisi! between thè neuropodial and notopodial tufts of
28
GIACOMO BRACCHI & ANNA ALESSANDRELLO
each parapodium nor to hypothesize whether thè dorsal or ventral surfaces of thè examined specimens are exposed. Each tuft is composed of several thread-like chaetae, measuring from 2 mm (MSNM Ì23085) to 6 mm (MSNM Ì23086) in length, and of a smaller number of stronger and thicker chaetae, which may be aciculae, preserved in MSNM Ì23087, MSNM Ì12397 and MSNM Ì24397. The shorter chaetae are those of thè anterior and posterior metameres, while thè longer ones are those of thè centrai metameres. Most of thè chaetae are directed backward but thè number of elements that equip each tuft may not be recognized due to thè poor quality of preservation.
Remarks : thè generai body morphology allows thè examined material to be related to family Aphroditidae, while thè arrangement of thè chaetae in two tufts on thè sides of each metamere suggests a relationship with thè genus Paleoaphrodite. The Triassic genus Homaphrodite is slightly different from Paleoaphrodite in thè generai morphology of thè body and in thè architecture of thè aciculae. The shape of H. speciosa resembles closely that of a typical worm-like metazoan and thè specimen described by Gali & Grauvogel (1966) seems to pos- sess only thin chaetae: on thè contrary, thè body of P. libanotica n. sp. has thè typical shape of a scale-worm and it is equipped with two chaetal types. However, thè two genera are hardly comparable as H. speciosa has thè antennae and elytrae preserved and these are not evident in P. libanotica n. sp.
The studied specimens show a strong similarity to two of thè species described to date under Paleoaphrodite : thè Triassic species P. raetica and thè Jurassic species P. gallica (Alessandrello & Teruzzi, 1986b; Alessandrello et al., 2004). P. libanotica n. sp. is similar to P. raetica in thè oval/elliptical shape of thè body but it differs from P. raetica in thè strong similarity between thè notopodial and neuropodial chaetae. This means that in P libanotica n. sp. each setal tuft includes both thread-like chateae and strong chaetae, while in P raetica a tuft is formed of thread-like chaetae only while thè other tuft includes both thin and strong chaetae. In P. gallica thè two tufts of each parapodium are composed of both strong and thin chaetae as in P. libanotica n. sp. but thè body is fusiform.
Superfamily Glyceroidea Grube, 1 850 Family Goniadidae Kingberg, 1866 Genus Ferragutia n. gen.
Diagnosis : as thè type species.
Derivatio nominis : dedicated to Marco Ferraguti, who kindly donated one of thè specimens among thè examined material.
Type species: Ferragutia cenomaniana n. sp.
Ferragutia cenomaniana n. sp.
PI. XIV
Diagnosis : pharyngeal apparatus armoured with a micrognaths circlet and two rows of chevrons. Macrog- naths absent. Presence of longitudinal muscle bundles.
Derivatio nominis : referred to thè geological age of thè Haqel outcrop.
Geological age : Cenomanian.
Type locality : Haqel.
Material : 14 specimens. MSNM Ì8340 (holotype), MSNM il 2423, MSNM il 2426 (part and counterpart) and MSNM Ì25439 (paratypes), MSNM Ì9203, MSNM Ì9357, MSNM il 24 10 (part and counterpart) and MSNM il 3291 (part and counterpart), MSNM il 3306 I, MSNM il 3306 II, MSNM il 2425 (part and counterpart), MSNM il 6295, MSNM i 1 95 1 2, MSNM Ì24961 and MSNM Ì24973.
Description : thè presence of some whole-preserved specimens allowed thè body length to be measured, from 2.6 mm (MSNM Ì9357) to 54 mm (MSNM il 24 10 and MSNM il 3291), and thè maximum width, from 0.3 mm (MSNM Ì24973) to 3.5 mm (MSNM Ì12423). The width of thè body is not Constant but it seems to decrease from thè caudal extremity to thè cephalic one.
Most of thè examined specimens preserve both micro¬ gnaths and chevrons, while few specimens preserve thè chevrons only (MSNM Ì9203, MSNM il 24 10, MSNM i 1 329 1 and MSNM i 1 95 1 2). Sometimes only a single row of chevrons is preserved (MSNM Ì9203, MSNM Ì24973). The micrognaths and chevrons of thè examined specimens are preserved as small, black, shiny structures probably stili composed of thè originai heavily sclerotinized mate¬ rial enriched with metal-ions. MSNM il 2423 lacks both micrognaths and chevrons but can be reasonably assigned to thè goniadid here described because of thè presence of some longitudinal muscle bundles. In thè whole-pre¬ served specimens identical soft structures are particularly evident in thè posterior part of thè body. Also thè rela¬ tive dimensions of MSNM i 12423, 53 mm in length and 3.5 mm in width at thè mid-length, suggest thè affinites with thè other specimens described here. The presence of longitudinal muscle bundles in specimen MSNM Ì24961, which has only a short length of thè posterior part of thè body preserved, allows comparisons with F. cenomaniana n. gen. n. sp. Near thè more slender extremity of this short section a low-preserved eunicemorph stomodaeal appa¬ ratus is preserved: it may represent evidence of predation even if it is difficult to distinguish whether thè goniadid body and thè eunicemorph apparatus are simply superim- posed.
The poor quality of preservation of many examined ! specimens contributed to thè distortion (MSNM Ì9357 and MSNM il 2426), splitting (MSNM Ì8340, MSNM : Ì9203 and MSNM Ì25439), disturbale (MSNM il 2425) - or/and interference (MSNM il 24 10 and MSNM il 3291) of thè rows of chevrons. However, well-preserved rows of chevrons are evident in MSNM il 95 12, where a single row of 20 elements is preserved, in MSNM Ì24973, where a single row (0.7 mm long) of 30 elements is preserved at a distance of 0.5 mm from thè micrognaths, and in MSNM i 1 3306 II, where both rows are preserved at a distance of 1 mm from thè micrognaths. The largest chevrons are those of thè centrai part of each row, while thè smallest are those of thè extremities. The micrognaths are also often disturbed or disrupted so that they form a mass in which their borders and shape may no longer be distinguished.
Only a few specimens preserve some imprints of , aciculae that measure from 0.3 mm (MSNM Ì24973) to 0.7 mm (MSNM il 24 10 and MSNM il 3291) in length. In MSNM il 2423 thè aciculae do not appear on both sides of thè body as in thè other specimens that show a dorsoventral preservation however, particularly in thè posterior sector, their arrangement seems to support a
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON 29
; lateral compression during diagenesis. The poor quality I of preservation of thè examined material did not facili- ; tate an investigation of how thè aciculae provide each parapodium and in turn provide information regarding where thè parapodia are uniramous and where biramous: furthermore, in modem goniadids thè parapodia are uniramous anteriorly, biramous posteriorly (Fauchald,
; 1977).
In addition to thè longitudinal muscle bundles (MSNM il 2401, MSNM i 1 2423, MSNM Ì13291, MSNM Ì19512 and MSNM Ì24961) thè preserved soft parts include thè body outline and dark traces of thè alimentary canal, particularly evident in thè posterior part of thè body. The trunk segmentation is hardly recognizable.
Remarks : a pharyngeal armature consisting of micrognaths and/or chevrons is typical of thè gonia- did polychaetes. The stomodaeum of thè goniadids is an axial eversible pharynx (proboscis) armoured with jaws around thè terminal aperture and, in some taxa, also with chevrons toward thè base. The jaws consist of a circlet of micrognaths sometimes added to some paired macrognaths that appear as larger and denticulate j structures. The chevrons support thè proboscis and are usually arranged along two longitudinal rows (Hartman, i 1950; Fauchald & Rouse, 1997). However, among thè
examined specimens evidence of an everted pharynx has not been found.
Among thè modem goniadids three genera possess chevrons (Hartman, 1950; Fauchald, 1977; Fauchald & Rouse, 1977) in thè same way as Ferragutia n. gen.: Goniada , Goniadella Hartman, 1950 and Progoniada Hartman, 1965. Neverthless, thè lack of macrognaths in all thè examined specimens, can hardly be considered an artifact of fossilization but rather as due to a prospective primitive organization (Hartman, 1950), which discrimi- nates Ferragutia n. gen. from thè modem representatives of this family and from thè Missisipian fossil species described by Schram (1979) as Carbosesostris meg- aliphagon. The lack of macrognaths has already been observed in 32 specimens of thè Carboniferous Pieckonia heìenae, which may be distinguished from Ferragutia cenomaniana n. sp. as it also lacks chevrons (Thompson, 1979).
Furthermore, thè pharyngeal armature of Ferragutia n. gen. seems strongly sclerotinized and strengthened by metal-ions in thè same way as thè living genera Glycinde Miiller, 1858 and Bathyglycinde Fauchald, 1972, whose pharynxes are characterized, in contrast to Ferragutia n. gen., by thè absence of thè chevrons (Hartman, 1950; Fauchald, 1977).
PALEOBIOLOGY AND PALEOECOLOGY
Feeding habits
A lot of thè considered specimens preserve a strong pharyngeal apparatus which, in some cases may sug- gest a predatory habit. However, among thè modern polychaetes that possess a stomodaeum armoured with jaws and maxillae, some taxa are omnivorous (Kay & Brafield, 1973) or detritivorous (Sanders, 1956; Banse & Hobson, 1968; Goerke, 1971). The carnivorous polychaetes predate crustaceans, molluscs, fìsh, nema- todes or other worms, but also less active preys, such as hydrozoans, bryozoans and poriferans (Mileikovsky, 1962; Hamond, 1969). Among thè examined material several specimens preserve traces of thè digestive tube as homogeneous, rust coloured thin bands. The pres¬ ervation of these traces has probably been assisted by thè presence of alimentary remains inside thè digestive ube at thè time of death, however, thè food source is never recognizable. Nevertheless, a predatory habit for thè aphroditid Paleoaphrodite libanotica n. sp. and for thè goniadid Ferragutia cenomaniana n. gen. n. sp. are hypothesized here. The undetermined MSNM il 2402 (Fig. 3) preserves a nematode buccal apparatus along thè dark trace of thè alimentary canal, thè aphroditid specimen MSNM Ì24937 (Fig. 4) has 2-3 vertebrae of a tiny fìsh preserved along thè reddish trace of thè ali¬ mentary canal, thè goniadid specimen MSNM Ì24961 preserves an eunicemorph pharyngeal armature in thè posterior part of thè body. The modern goniadids are actually carnivorous and they use their eversible proboscis crowned with jaws to capture prey (Hart- ' man, 1950): only a few species use their jaws to grasp detritus particles. The modern aphroditids generally live in and on soft muddy bottoms where they slowly bulldoze their way through thè sediment in search of
organic detritus and small infauna to feed on. Although other living aphroditids tend to be carnivorous on small invertebrates and vertebrates rather than being primarily detritivorous like thè sand-dwelling species (Hutchings & McRae, 1993).
Fig. 3 - MSNM il 2402, undetermined specimen preserving possible traces of predation.
30
GIACOMO BRACCHI & ANNA ALESSANDRELLO
Reproduction
Among thè living polychaetes some particular repro- ductive phoenomena, such as epitoky and stolonization, are known (Giangrande, 1997). The epitokous specimens are pelagic reproductive specimens that originate from thè atokous specimens, or rather from non-reproductive individuals that undergo a direct, complete and profound trasformation, as happens in some nereidids. In other cases thè atokous specimens are simply involved in thè trasformation and separation of thè posterior part of thè body (schizogamy), as happens in some syllids and eunicids. The epitokous metameres appear profoundly modified with respect to thè others so that thè body of thè
Fig. 4 - Paleoaphrodite libanotica n. sp., MSNM Ì24937, preserving possible traces of predation.
worm seems to be divided in two morphologically distinct regions.
In thè same way of thè epitochy, thè stolonization is an sexual reproductive process that implies thè for- mation of some stolons at thè caudal extremity of thè specimens involved: through a rapid growth and their consequent separation, thè stolons become new indi¬ viduals.
Among thè examined material, thè indeterminate specimen MSNM Ì25433 (Fig. 5) shows profound mor- phological and morphometrical differences between thè anterior and posterior metameres: this may be evidence of epitochy or stolonization but thè poor quality of preserva- tion did not allow further information to be obtained.
Fig. 5 - MSNM Ì25433, undetermined specimen showing possible traces of epitcchy/stolonization.
DISCUSSION AND CONCLUSIONS
With 379 specimens thè Lebanese collection of thè fossil free-living polychaetes held by thè Museo Civico di Storia Naturale di Milano represents thè richest and most varied polychaete paleofauna known within thè Mesozoic and, after thè Field Museum (Chicago, U.S.A.) collection from thè Carboniferous of Mazon Creek (Thompson, 1979), one of thè most renowned and numerous in thè world. A systematic definition at species level has been possible for 114 specimens: 2 specimens have been already described by Alessandrello & Teruzzi (1986), 112 (110 from Haqel, 2 from Hadjula) have been described in this work. To date 2 orders (Eunicida and Phyllodocida), 6 families (Aphroditidae, Didonidae n. fam., Eunicidae,
Goniadidae, Lumbrineridae and Oenonidae), 7 genera ( Bidone n. gen., Eunicites, Ferragutia n. gen., Lumbri- conerites, Paleoaphrodite , Phoeniciarabella n. gen. and Teruzzia n. gen.) and 17 species ( Didone pulcherrima n. gen. n. sp., Eunicites diopatroides n. sp., Eunicites falca- tus n. sp., Eunicites joinvillei n. sp., Eunicites mariacristi- nae n. sp., Eunicites phoenicius, Ferragutia cenomaniana \ n. gen. n. sp., Lumbriconereites garassinoi n. sp., Lum- briconereites hadjulae n. sp., Paleoaphrodite libanotica n. sp., Phoeniciarabella caesaris n. gen. n. sp., Phoeni¬ ciarabella orensanzi n. gen. n. sp., Phoeniciarabella pin- nulata n. gen. n. sp., Teruzzia gryphoeides n. gen. n. sp., Teruzzia pezzolii n. gen. n. sp., Teruzzia pusilla n. gen. n.
PALEODIVERSITY OF THE FREE-LIVTNG POLYCHAETES (ANNEL1DA, POLYCHAETA) AND DESCR1PTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON 3 J
sp. and Teruzzia sagittifera n. gen. n. sp.) of polychaetous annelids have been recorded from thè Cenomanian fos- siliferous levels of Lebanon. A systematic definition to species level was not possible for 265 specimens (Fig.
6) : 226 stili remain undetermined, due to thè absence of diagnostic features related to thè architecture of thè pharyngeal armature or to thè soft tissues that are rarely preserved in fossils, 31 have been classified to generic level (1 Lumbriconereites, 4 Eunicites and 26 Teruzzia n. gen.) while only 8 were identified to family level (8 Lumbrineridae).
Among thè 363 specimens from Haqel there are (Fig.
7) 212 undetermined specimens, 106 Lumbrineridae, 16 Eunicidae, 14 Goniadidae, 6 Aphroditidae, 5 Didonidae n. fam. and 4 Oenonidae: thè most represented genera are thè lumbrinerid Teruzzia n. gen. (82) and Lumbriconerites (16), followed by thè eunicid Eunicites (16), thè goniadid Ferragutia n. gen. (14), thè aphroditid Paleoaphrodite (6), thè didonid Didone n. gen. (5) and thè oenonid Phoe- niciarabella n. gen. (4). Regarding thè Haqel outcrop, thè species that include thè greatest number of specimens are Teruzzia pezzolii n. gen. n. sp. (52) and Lumbriconerites
hadjulae n. sp. (14), followed by Ferragutia cenomani- ana n. gen. n. sp. (14), Paleoaphrodite libanotica n. sp. (6), Eunicites joinvillei n. sp. (5), Didone pulcherrima n. gen. n. sp. (5), Eunicites diopatroides n. sp. (3), Eunicites phoenicius (2), Teruzzia sagittifera n. gen. n. sp. (2) and Phoeniciarabella caesaris n. gen. n. sp. (2). From thè same outcrop only thè holotype has been described for Eunicites falcatus n. sp., Eunicites mariacristinae n. sp., Lumbriconereites garassinoi n. sp., Phoeniciarabella orensanzi n. gen. n. sp., Phoeniciarabella pinnulata n. gen. n. sp., Teruzzia gryphoeides n. gen. n. sp. and Ter¬ uzzia pusilla n. gen. n. sp.
With regard to thè outcrops of Hadjula and Al- Namoura there are few data on thè polychaete paleodiver- sity as only 15 specimens for thè former and 1 specimen for thè latter have been collected from these sites to date. Futhermore, among thè 15 specimens from Hadjula, a systematic definition to thè species level has been pos¬ sible only for two: they belong to Lumbriconereites had¬ julae n. sp.
To date an investigation of thè relative abundance of thè polychaete families from a single paleobasin has been
18,4%
B Nephtyidae
■ Aphroditidae Fossundecimidae Eunicidae
■ Goniadidae
■ Amphinomidae
■ Serpuliidae Phyllodocidae
■ Flabelligeridae
■ Hesionidae Opheliidae
■ Palmyridae
Fig. 6 - Distribution of thè examined material among undetermined specimens, determined to species level, generic level or to family level.
■ undetermined
■ determinated sp. level determinated gen. level
■ determinated fam. level
Fig. 7 - Distribuiton of thè examined material from Haqel among undetermined specimens, Eunicidae, Lumbrineridae, Oenonidae, Didonidae, Aphroditidae and Goniadidae.
32
GIACOMO BRACCHI & ANNA ALESSANDRELLO
possible only for thè Essex Carboniferous biota where at least 1085 polychaete specimens assigned to 12 families have been discovered (Thompson & Johnson, 1977; Thompson, 1979; Hay, 2002). The first difference that can be pointed out through thè comparison of thè polychaete fauna from Haqel and that from Essex is thè presence of sedentary polychaetes in thè latter: they represent about 3 % of thè total specimens and they have been assigned, in order of abundance, to thè flabelligerids, serpulids and opheliids. With regard to thè free-living polycha¬ etes, thè Essex fauna is mostly comprised of, in order of abundance, nephtyids, aphroditids, fossundecimids and eunicids, followed by small percentages of amphinomids, goniadids, phyllodocids, hesionids and palmyrids (Fig.
8). The differences between thè polychaete fauna from Haqel and that from Mazon Creek are probably related to thè different paleoenvironments of these outcrops. Haqel was a small, deep paleobasin characterized by restriction of water circulation, stagnation, hypersalinity and oxygen depletion. On thè contrary thè Francis Creek Member from which most of thè specimens described by Thompson (1979) and Hay (2002) were collected represents a delta complex influenced by sea-level and salinity fluctuations. There were distributary channels, interdistributary bays and a proximal prodelta as well as transitions from marine to estuarine and terrestrial facies (Schellenberg, 2002).
29%
■ undetermined
■ Eunicidae Lumbrineridae
■ Oenonidae Didonidae
■ Aphroditidae
■ Goniadidae
Fig. 8 - Systematic distribution of thè polychaete fauna ffom Mazon Creek according to Thompson (1979) and Hay (2002).
Acknowledgements
We wish to thank Fabio Marco Dalla Vecchia (Museo Paleontologico Cittadino, Monfalcone), Maria Cristina Gambi (Laboratorio di Ecologia del Benthos, Stazione Zoologica “Anton Dohm”, Napoli), Kristian Fauchald (National Museum of Naturai History, Smithsonian Insti- tution, Washington), Alessandro Garassino and Giorgio
Teruzzi (Dipartimento di Paleontologia, Museo Civico di Storia Naturale di Milano), for their useful advices during this work, Roberto Appiani and Franco Nodo (Museo Civico di Storia Naturale di Milano), for thè iconographic material, Kathleen Histon, for thè careful review of thè English text.
PALEODIVERSITY OF THE FREE-LIVTNG POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRJPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
1)1
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■
i
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Giacomo Bracchi - Museo Civico di Storia Naturale di Milano, Corso Venezia 55, 20121 Milano, Italy
e-mail: mojak@inwind.it
Anna Alessandrello - Museo Civico di Storia Naturale di Milano, Corso Venezia 55, 20121 Milano, Italy
e-mail: anna.alessandrello@comune.milano.it
Paleodiversity of thè free-living polychaetes (Annelida, Polychaeta) and description of new taxa from thè Upper Cretaceous
Lagerstàtten of Haque, Hadjula and Al-Namoura
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXII - Fascicolo III
40
GIACOMO BRACCHI & ANNA ALESSANDRELLO
Appendix - Complete listing of thè examined specimens
The specimens marked with are from Hadjula, those marked with from Al-Namoura, all thè others from Haqel.
MSNM vi 1681 |
und. |
und. |
und. |
und. |
|
MSNM vii 6811 |
und. |
und. |
und. |
und. |
|
MSNM vi 168III |
und. |
und. |
und. |
und. |
|
MSNM vll68IV |
und. |
und. |
und. |
und. |
|
MSNM vi 187 |
und. |
und. |
und. |
und. |
|
MSNM v2148 |
und. |
und. |
und. |
und. |
|
MSNM v3703 |
und. |
und. |
und. |
und. |
|
MSNM Ì7843 |
Eunicida |
Eunicidae |
Eunicites |
phoenicius |
holotype |
MSNM Ì7844 |
Eunicida |
Eunicidae |
Eunicites |
phoenicius |
paratype |
MSNM Ì8338 |
und. |
und. |
und. |
und. |
|
MSNM Ì8339 |
und. |
und. |
und. |
und. |
|
MSNM Ì8340 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
holotype |
MSNM Ì8342 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì8343 |
und. |
und. |
und. |
und. |
|
MSNM Ì8344 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì8345 |
und. |
und. |
und. |
und. |
|
MSNM Ì8346 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì8347 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì8348 |
und. |
und. |
und. |
und. |
|
MSNM Ì8349 |
und. |
und. |
und. |
und. |
|
MSNM Ì8350 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì8378 |
und. |
und. |
und. |
und. |
|
MSNM Ì8379 |
Eunicida |
Lumbrineridae |
Teruzzia |
pusilla |
holotype |
MSNM Ì8380 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì8381 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì8382 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i8383(A) + MSNM i8387(B) |
und. |
und. |
und. |
und. |
|
MSNM Ì8384 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì8385 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì8386 |
und. |
und. |
und. |
und. |
|
MSNM Ì8388 |
und. |
und. |
und. |
und. |
|
MSNM Ì8389 |
und. |
und. |
und. |
und. |
|
MSNM Ì8390 |
und. |
und. |
und. |
und. |
|
MSNM Ì8391 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì8472 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì8473 |
und. |
und. |
und. |
und. |
|
MSNM Ì8474 |
und. |
und. |
und. |
und. |
|
MSNM Ì8475 |
und. |
und. |
und. |
und. |
|
MSNM Ì8476 |
Eunicida |
Lumbrineridae |
Teruzzia |
sagittifera |
paratype |
MSNM Ì8479 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì8480 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì8481 |
und. |
und. |
und. |
und. |
|
MSNM Ì8482 |
und. |
und. |
und. |
und. |
|
MSNM Ì8483 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCFIAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
41
MSNM Ì8487 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM 18488 |
Eunicida |
Eunicidae |
Eunicites |
falcatus |
holotype |
MSNM Ì8489 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì8490 |
Eunicida |
Eunicidae |
Eunicites |
mariacristinae |
holotype |
MSNM i9 1 78 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i9 1 79 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 80 |
und. |
und. |
und. |
und. |
|
MSNM Ì9182I |
und. |
und. |
und. |
und. |
|
MSNM Ì9182II |
und. |
und. |
und. |
und. |
|
MSNM i9 1 83 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 84 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i9 1 85 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 86 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 87 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 90 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM i9 1 92 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM i9 1 93 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i9 1 94 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM i9 1 95 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM i9 1 96 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 97 |
und. |
und. |
und. |
und. |
|
MSNM i9 1 99 |
und. |
und. |
und. |
und. |
|
MSNM 9200 |
und. |
und. |
und. |
und. |
|
MSNM Ì9201 |
und. |
und. |
und. |
und. |
|
MSNM Ì9202 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì9203 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM 92061 |
und. |
und. |
und. |
und. |
|
MSNM 9206II |
und. |
und. |
und. |
und. |
|
MSNM Ì9207 |
und. |
und. |
und. |
und. |
|
MSNM Ì9208 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì9209 |
und. |
und. |
und. |
und. |
|
MSNM Ì9210 |
und. |
und. |
und. |
und. |
|
MSNM Ì9211 |
und. |
und. |
und. |
und. |
|
MSNM Ì9212 |
und. |
und. |
und. |
und. |
|
MSNM Ì9213 |
und. |
und. |
und. |
und. |
|
MSNM 19214 |
und. |
und. |
und. |
und. |
|
MSNM i9355(A)+MSNM i20618(B) |
Eunicida |
Lumbrineridae |
Lumbriconereites |
garassinoi |
holotype |
MSNM Ì9356 |
und. |
und. |
und. |
und. |
|
MSNM Ì9357 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM Ì9258 |
und. |
und. |
und. |
und. |
|
MSNM Ì9359 (A+B) |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì9360 |
und. |
und. |
und. |
und. |
|
MSNM Ì9361 |
und. |
und. |
und. |
und. |
|
MSNM Ì9362 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
**MSNM Ì9363 |
und. |
und. |
und. |
und. |
|
MSNM Ì9367 |
und. |
und. |
und. |
und. |
42
GIACOMO BRACCHI & ANNA ALESSANDRELLO
MSNM Ì9369 |
und. |
und. |
und. |
und. |
|
MSNM Ì9370 |
und. |
und. |
und. |
und. |
|
MSNM il 0402 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
paratype |
MSNM il 0762 |
und. |
und. |
und. |
und. |
|
MSNM il 0763 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM il 0968 |
und. |
und. |
und. |
und. |
|
MSNM il 2396 |
und. |
und. |
und. |
und. |
|
MSNM i 1 2397 |
Phyllodocida |
Aphroditidae |
Paleoaphrodite |
libanotica |
|
MSNM i 1 2398 |
und. |
und. |
und. |
und. |
|
MSNM il 2400 |
und. |
und. |
und. |
und. |
|
MSNM Ì12401 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 2402 |
und. |
und. |
und. |
und. |
|
MSNM il 2403 |
und. |
und. |
und. |
und. |
|
MSNM il 2404 |
und. |
und. |
und. |
und. |
|
MSNM i 1 2405 |
und. |
und. |
und. |
und. |
|
MSNM il 2406 |
und. |
und. |
und. |
und. |
|
MSNM il 2407 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i 1 2408( A)+MSNM il2412(B) |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 2409 |
und. |
und. |
und. |
und. |
|
MSNM i 1 24 1 0(A)+MSNM il3291(B) |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM il 24 11 |
Eunicida |
Oenonidae |
Phoeniciarabella |
pinnulata |
holotype |
MSNM il 24 13 |
und. |
und. |
und. |
und. |
|
MSNM i 1 24 1 4 |
Eunicida |
Eunicidae |
Eunicites |
joinvillei |
paratype |
MSNM i 1 24 1 5(A)+MSNM il3305(B) |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i 1 24 1 7 |
und. |
und. |
und. |
und. |
|
MSNM il 24 19 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 2420 |
Eunicida |
Eunicidae |
Eunicites |
joinvillei |
holotype |
MSNM il 2421 |
und. |
und. |
und. |
und. |
|
MSNM il 2422 |
und. |
und. |
und. |
und. |
|
MSNM il 2423 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
paratype |
MSNM il 2424 (A+B) |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
holotype |
MSNM il 2425 (A+B) |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM il 2426 (A+B) |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
paratype |
MSNM il 2626 |
und. |
und. |
und. |
und. |
|
MSNM i 1 2628 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM il 2629 |
und. |
und. |
und. |
und. |
|
MSNM i 1 2630 |
und. |
und. |
und. |
und. |
|
MSNM il 2631 |
und. |
und. |
und. |
und. |
|
MSNM il 2632 (A+B) |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
paratype |
MSNM il 2936 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM il 3283 |
Eunicida |
Oenonidae |
Phoeniciarabella |
orensanzi |
holotype |
MSNM il 3284 |
Eunicida |
Eunicidae |
Teruzzia |
pezzolii |
|
MSNM i 1 3286 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM i 1 3287 |
und. |
und. |
und. |
und. |
|
MSNM il 3288 (A+B) |
Eunicida |
Eunicidae |
Eunicites |
diopatroides |
paratype |
MSNM i 1 3289 |
und. |
und. |
und. |
und. |
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCFLAETA) AND DESCR1PTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
MSNM il 3290 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
holotype |
MSNM il 3292 |
und. |
und. |
und. |
und. |
|
MSNM il 3293 |
und. |
und. |
und. |
und. |
|
MSNM il 3294 |
und. |
und. |
und. |
und. |
|
MSNM il 3295 |
und. |
und. |
und. |
und. |
|
MSNM i 1 3296 |
und. |
und. |
und. |
und. |
|
MSNM il 3297 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 3298 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 3299 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM i 1 3301 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 3302 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 3303 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM il 3304 |
Eunicida |
Eunicidae |
Eunicites |
joinvillei |
paratype |
MSNM il 33061 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM i 1 3306II |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM il 3307 |
und. |
und. |
und. |
und. |
|
MSNM i 13308 |
und. |
und. |
und. |
und. |
|
MSNM il 3309 |
und. |
und. |
und. |
und. |
|
MSNM il 331 1 |
Eunicida |
Eunicidae |
Eunicites |
und. |
|
MSNM i 133 12 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM il3606(A) + MSNM il6303(B) |
und. |
und. |
und. |
und. |
|
MSNM il 62751 |
und. |
und. |
und. |
und. |
|
MSNM i 1 6275II |
und. |
und. |
und. |
und. |
|
MSNM Ì16276 |
und. |
und. |
und. |
und. |
|
MSNM il 6278 |
und. |
und. |
und. |
und. |
|
MSNM il 6279 |
Euncida |
Lumbrineridae |
und. |
und. |
|
MSNM il 6282 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM il 6287 |
und. |
und. |
und. |
und. |
|
MSNM il 6288 |
Phyllodocida |
Aphroditidae |
Paleoaphrodite |
libano tic a |
holotype |
MSNM il 6291 |
und. |
und. |
und. |
und. |
|
MSNM il 6293 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM il 6294 |
und. |
und. |
und. |
und. |
|
MSNM il 6295 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM il 6296 |
und. |
und. |
und. |
und. |
|
MSNM il 6299 |
und. |
und. |
und. |
und. |
|
MSNM i 1 6308 |
und. |
und. |
und. |
und. |
|
MSNM il 95 12 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM Ì20654 |
und. |
und. |
und. |
und. |
|
MSNM Ì20593 |
Eunicida |
Eunicidae |
Eunicites |
und. |
|
MSNM Ì20595 |
und. |
und. |
und. |
und. |
|
MSNM Ì20597 |
und. |
und. |
und. |
und. |
|
MSNM Ì20599 |
und. |
und. |
und. |
und. |
|
MSNM Ì20600 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20601 |
und. |
und. |
und. |
und. |
|
MSNM Ì20603I |
und. |
und. |
und. |
und. |
|
MSNM Ì20603II |
und. |
und. |
und. |
und. |
44
GIACOMO BRACCHI & ANNA ALESSANDRELLO
MSNM Ì20608 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20609 |
und. |
und. |
und. |
und. |
|
MSNM Ì20610 (A+B) |
Eunicida |
Oenonidae |
Phoeniciarabella |
caesaris |
holotype |
MSNM Ì20611 |
Euncida |
Lumbrineridae |
Teruzzia |
pezzolii |
paratype |
MSNM Ì20612 |
Eunicida |
Eunicidae |
Eunicites |
joinvillei |
|
MSNM Ì20613 |
Eunicida |
Eunicidae |
Eunicites |
diopatroides |
paratype |
MSNM Ì20614 |
Eunicida |
Eunicidae |
Lumbriconereites |
hadjulae |
paratype |
MSNM Ì20615 |
und. |
und. |
und. |
und. |
|
MSNM Ì20616 |
und. |
und. |
und. |
und. |
|
MSNM Ì20617 (A+B) |
Eunicida |
Didonidae |
Diclone |
pulcherrima |
holotype |
MSNM Ì20619 |
Eunicida |
Didonidae |
Bidone |
pulcherrima |
|
MSNM Ì20620 |
und. |
und. |
und. |
und. |
|
MSNM Ì20621 |
und. |
und. |
und. |
und. |
|
MSNM Ì20623 |
Eunicida |
Eunicidae |
Lumbriconereites |
hadjulae |
|
MSNM Ì20624 |
Eunicida |
Didonidae |
Didone |
pulcherrima |
paratype |
MSNM Ì20625 |
und. |
und. |
und. |
und. |
|
MSNM Ì20627 |
und. |
und. |
und. |
und. |
|
MSNM Ì20628 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20629 |
und. |
und. |
und. |
und. |
|
MSNM Ì20630 |
und. |
und. |
und. |
und. |
|
MSNM Ì20632 |
und. |
und. |
und. |
und. |
|
MSNM Ì20633 |
und. |
und. |
und. |
und. |
|
MSNM Ì20638 |
und. |
und. |
und. |
und. |
|
MSNM Ì20639 |
und. |
und. |
und. |
und. |
|
MSNM Ì20640 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20641 |
und. |
und. |
und. |
und. |
|
MSNM Ì20642 |
und. |
und. |
und. |
und. |
|
MSNM Ì20643 |
und. |
und. |
und. |
und. |
|
MSNM Ì20644 |
und. |
und. |
und. |
und. |
|
MSNM Ì20645 |
und. |
und. |
und. |
und. |
|
MSNM Ì20646 |
und. |
und. |
und. |
und. |
|
MSNM Ì20647 |
und. |
und. |
und. |
und. |
|
MSNM Ì20648 |
und. |
und. |
und. |
und. |
|
MSNM Ì20649 |
und. |
und. |
und. |
und. |
|
MSNM Ì20651 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20653 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20654 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20656 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20657 |
und. |
und. |
und. |
und. |
|
MSNM Ì20658 |
und. |
und. |
und. |
und. |
|
MSNM Ì20659 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì20661 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì20662 |
und. |
und. |
und. |
und. |
|
MSNM Ì20663 |
und. |
und. |
und. |
und. |
|
MSNM Ì22829 |
und. |
und. |
und. |
und. |
|
MSNM Ì22830 |
und. |
und. |
und. |
und. |
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
MSNM Ì22831 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì22832 |
und. |
und. |
und. |
und. |
|
MSNM Ì22833 |
und. |
und. |
und. |
und. |
|
MSNM Ì22835 |
und. |
und. |
und. |
und. |
|
MSNM Ì22836 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzo Hi |
|
MSNM Ì22837 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì22838 |
und. |
und. |
und. |
und. |
|
MSNM Ì22839 |
und. |
und. |
und. |
und. |
|
MSNM Ì22840 |
und. |
und. |
und. |
und. |
|
MSNM Ì23046 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì23048 |
und. |
und. |
und. |
und. |
|
MSNM Ì23051 |
und. |
und. |
und. |
und. |
|
MSNM Ì23053 |
und. |
und. |
und. |
und. |
|
MSNM Ì23056 |
und. |
und. |
und. |
und. |
|
MSNM Ì23058 |
und. |
und. |
und. |
und. |
|
MSNM Ì23059 |
und. |
und. |
und. |
und. |
|
MSNM Ì23083 |
und. |
und. |
und. |
und. |
|
MSNM Ì23084 |
Eunicida |
Eunicidae |
Eunicites |
diopatroides |
holotype |
MSNM Ì23085 |
Phyllodocida |
Aphroditidae |
Paleoaphrodite |
libanotica |
paratype |
MSNM Ì23086 |
Phyllodocida |
Aphroditidae |
Paleoaphrodite |
libanotica |
paratype |
MSNM Ì23087 |
Phyllodocida |
Aphroditidae |
Paleoaphrodite |
libanotica |
|
MSNM Ì23414 |
und. |
und. |
und. |
und. |
|
MSNM Ì23415 |
Eunicida |
Didonidae |
Didone |
pulcherrima |
|
MSNM Ì23416 |
und. |
und. |
und. |
und. |
|
MSNM Ì23417I |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì23417II |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì23419I |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì23419II |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì23421 |
Eunicida |
Oenonidae |
Phoeniciarabelìa |
caesaris |
paratype |
MSNM Ì23424 |
und. |
und. |
und. |
und. |
|
MSNM Ì23425 |
und. |
und. |
und. |
und. |
|
MSNM Ì23426 |
und. |
und. |
und. |
und. |
|
MSNM Ì23427 |
und. |
und. |
und. |
und. |
|
MSNM Ì23428 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì23429 |
und. |
und. |
und. |
und. |
|
MSNM Ì23436 |
Eunicida |
Lumbrineridae |
Teruzzia |
s agi t tiferà |
holotype |
MSNM Ì23440 |
und. |
und. |
und. |
und. |
|
MSNM Ì23441 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì23443 |
und. |
und. |
und. |
und. |
|
MSNM Ì23447 |
und. |
und. |
und. |
und. |
|
MSNM Ì23448 |
und. |
und. |
und. |
und. |
|
MSNM Ì23450 |
und. |
und. |
und. |
und. |
|
MSNM Ì23453 |
und. |
und. |
und. |
und. |
|
MSNM Ì23454 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM Ì23455 |
und. |
und. |
und. |
und. |
|
MSNM Ì23457 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
46
GIACOMO BRACCHI & ANNA ALESSANDRELLO
MSNM Ì23458 |
und. |
und. |
und. |
und. |
|
MSNM Ì24812 |
Eunicida |
Eunicidae |
Eunicites |
und. |
|
MSNM Ì24813 |
und. |
und. |
und. |
und. |
|
MSNM Ì24814 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì24911 |
und. |
und. |
und. |
und. |
|
MSNM Ì24913 |
Eunicida |
Lumbrineridae |
und. |
und. |
|
MSNM Ì24915 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì24917 |
und. |
und. |
und. |
und. |
|
MSNM Ì24918 |
und. |
und. |
und. |
und. |
|
MSNM Ì24919 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì24926 (A+B) |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzo! ii |
|
MSNM Ì24930 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì24933 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì24935 |
und. |
und. |
und. |
und. |
|
MSNM Ì24936 (A+B)I |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì24936 (A+B)II |
und. |
und. |
und. |
und. |
|
MSNM Ì24937 |
Phyllodocida |
Aphroditidae |
Paleoaphrodite |
libanotica |
|
MSNM Ì24940 |
und. |
und. |
und. |
und. |
|
MSNM Ì24943 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
paratype |
MSNM Ì24944 |
und. |
und. |
und. |
und. |
|
MSNM Ì24945 |
und. |
und. |
und. |
und. |
|
MSNM Ì24946 |
und. |
und. |
und. |
und. |
|
MSNM Ì24961 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM Ì24965 |
und. |
und. |
und. |
und. |
|
MSNM Ì24964 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì24966 |
und. |
und. |
und. |
und. |
1 |
MSNM Ì24967 |
Eunicida |
Lumbrineridae |
Teruzzia |
gryphoeides |
holotype |
MSNM Ì24970 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì24972 |
und. |
und. |
und. |
und. |
|
MSNM Ì24973 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
|
MSNM Ì24976 |
und. |
und. |
und. |
und. |
|
MSNM Ì24977 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì24989 |
und. |
und. |
und. |
und. |
. |
MSNM Ì24990 |
und. |
und. |
und. |
und. |
|
MSNM Ì24991 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì24992 |
und. |
und. |
und. |
und. |
|
MSNM Ì24944 |
und. |
und. |
und. |
und. |
|
MSNM Ì25000 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì25001 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì25002 |
und. |
und. |
und. |
und. |
|
MSNM Ì25003 |
und. |
und. |
und. |
und. |
|
MSNM Ì25004 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzolii |
|
MSNM Ì25005 |
und. |
und. |
und. |
und. |
|
MSNM Ì25007 |
und. |
und. |
und. |
und. |
|
MSNM Ì25009 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì25011 |
und. |
und. |
und. |
und. |
PALEODIVERSITY OF THE FREE-LFVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
47
MSNM Ì25014 |
und. |
und. |
und. |
und. |
|
MSNM Ì25015 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì25019 |
und. |
und. |
und. |
und. |
|
MSNM Ì25020 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì25088 |
und. |
und. |
und. |
und. |
|
MSNM Ì25089 |
und. |
und. |
und. |
und. |
|
MSNM Ì25090 |
und. |
und. |
und. |
und. |
|
MSNM Ì25091 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì25092 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì25093 (A+B) |
und. |
und. |
und. |
und. |
|
*MSNM Ì25107 |
und. |
und. |
und. |
und. |
|
*MSNM Ì25108 |
und. |
und. |
und. |
und. |
|
*MSNM Ì25109 |
und. |
und. |
und. |
und. |
|
*MSNM Ì25110 |
und. |
und. |
und. |
und. |
|
*MSNM Ì25113 |
und. |
und. |
und. |
und. |
|
*MSNM Ì25115 (A+B) |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
*MSNM Ì25116 |
und. |
und. |
und. |
und. |
|
MSNM Ì25117 |
und. |
und. |
und. |
und. |
|
MSNM Ì25118 |
Eunicida |
Eunicidae |
Eunicites |
und. |
|
*MSNM Ì25119 |
und. |
und. |
und. |
und. |
|
*MSNM Ì25120 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì25433 |
und. |
und. |
und. |
und. |
|
MSNM Ì25434 |
und. |
und. |
und. |
und. |
|
MSNM Ì25435 |
und. |
und. |
und. |
und. |
|
MSNM Ì25436 |
und. |
und. |
und. |
und. |
|
MSNM Ì25437 |
und. |
und. |
und. |
und. |
|
MSNM Ì25438 |
Eunicida |
Lumbrineridae |
Teruzzia |
pezzo lii |
|
MSNM Ì25439 |
Phyllodocida |
Goniadidae |
Ferragutia |
cenomaniana |
paratype |
MSNM Ì25440 |
und. |
und. |
und. |
und. |
|
MSNM Ì25441 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì25442 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
*MSNM Ì25443 |
und. |
und. |
und. |
und. |
|
MSNM Ì25444 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì25445 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì25446 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
hadjulae |
|
MSNM Ì25447 |
und. |
und. |
und. |
und. |
|
MSNM Ì25448 |
und. |
und. |
und. |
und. |
|
MSNM Ì25449 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì25450 |
und. |
und. |
und. |
und. |
|
MSNM Ì25451 |
und. |
und. |
und. |
und. |
|
MSNM Ì25452 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì25453 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
MSNM Ì25454 |
Eunicida |
Lumbrineridae |
Teruzzia |
und. |
|
rMSNM Ì25455 (A+B) |
und. |
und. |
und. |
und. |
|
MSNM Ì25456 |
Eunicida |
Eunicidae |
Eunicites |
joinvillei |
|
MSNM Ì25457 |
Eunicida |
Didonidae |
Didone |
pulcherrima |
paratype |
48
GIACOMO BRACCHI & ANNA ALESSANDRELLO
MSNM Ì26326 |
und. |
und. |
und. |
und. |
|
MSNM Ì26327 |
und. |
und. |
und. |
und. |
|
MSNM Ì26328 |
und. |
und. |
und. |
und. |
|
MSNM Ì26329 |
und. |
und. |
und. |
und. |
|
MSNM Ì26330 |
Eunicida |
Lumbrineridae |
Lumbriconereites |
und. |
|
*MSNM Ì26332 |
und. |
und. |
und. |
und. |
|
MSNM Ì26333 |
und. |
und. |
und. |
und. |
|
*MSNM Ì26335 (A+B) |
und. |
und. |
und. |
und. |
|
*MSNM Ì26336 |
und. |
und. |
und. |
und. |
|
MSNM Ì26337 |
und. |
und. |
und. |
und. |
|
*MSNM Ì26338 |
und. |
und. |
und. |
und. |
|
MSNM Ì26339 |
und. |
und. |
und. |
und. |
|
MSNM Ì26340 |
und. |
und. |
und. |
und. |
C = maxillary carriers F = forceps J = jaws
PLATES I - XVI: LIST OF ABBREVI ATIONS
Japp = jaw apparatus M = maxilla
MP = mandibular processes
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
49
PI. I - Eunicites joinvillei n. sp., MSNM i 1 2420, holotype (A: x 4.8; B: x 4.5); Eunicites joinvillei n. sp., MSNM i 1 24 1 4, paratype (C: x 5.2; D: x 7.3).
50
GIACOMO BRACCHI & ANNA ALESSANDRELLO
A
B
C
PI. II - Eunicites falcatus n. sp., MSNM Ì8488, holotype (A: x 1 1.6; B: x 8.4); Eunicites diopatroides n. sp., MSNM Ì23084, holotype (C: x 7; D: x 7.4).
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
PI. Ili - Eunicites dìopatroides n. sp., MSNM Ì20613, paratype (A: x 10; B: x 10.3); Eunicites mariacristinae n. sp., MSNM Ì8490. holotype (C: x 100; D: x 120).
c D
PI. IV - Teruzzia pezzolii n. gen. n. sp., MSNM il 3290, holotype (A: x 93; B: x 94); Teruzzia pezzolii n. gen. n. sp., MSNM Ì2061 1 (C: x 23; D: x 28.5).
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
53
C
D
PI. V - Teruzzia sagittifera n. gen. n. sp., MSNM Ì23436, holotype (A: x 52.3; B: x 54.6); Teruzzia sagittifera n. gen. n. sp., MSNM Ì8476, paratype (C: x 47; D: x 54.6).
c
PI. VI - Te ruzzici pusilla n. gen. n. sp., MSNM Ì8379, holotype (A: x 10; B: x 128); Teruzzia gryphoeides n. gen. n. sp„ MSNM Ì24967 holotype (C: x 36.6; D: x 46.6).
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA. POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
PI. VII - Lumbriconereites hadjulae n. sp., MSNM il2424a, holotype, part, (A: x 9.5; B: x 16); Lumbriconereites hadjulae n. sp., MSNM Ì20614, paratype (C: x 8.8; D: x 16).
56
GIACOMO BRACCHI & ANNA ALESSANDRELLO
A
B
C
D
PI. Vili - Lumbrìconereites hadjulae n. sp., MSNM il2632a, holotype, part (A: x 10; B: x 17.5); Lumbriconereìtes hadjulae n. sp., MSNM i!2632b, holotype, counterpart (C: x 10; D: x 19.2).
PALEODIVERSITY OF THE FREE-LIVtNG POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
57
PI. IX - Lumbriconereites garassinoi n. sp., MSNM Ì9355, holotype (A: x 1 1 .3; B: x 11 .6); Phoeniciarabella pinnulata n. gen. n. sp MSNM i 1 24 1 1 , holotype (C: x 12; D: x 19).
58
GIACOMO BRACCHI & ANNA ALESSANDRELLO
c
D
PI. X - Phoeniciarabella orensanzi n. gen. n. sp., MSNM i 13283, holotype (A: x 0.7; B: x 360); Phoeniciar abella caesaris n. gen. n. sp., MSNM i 20610, holotype (C: x 10; D: x 18).
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCR1PTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
B
Japp
f
I
\
\
\
\
F \
V / I / t / l /
i c
il
il:
PI. XI - Phoeniciarabella caesaris n. gen. n. sp., MSNM Ì26421, paratype (A: x 3.6; B: x 36); Didone pulcherrima n. gen. n. sp., MSNM Ì20617, holotype (C: x 9.3; D: x 17).
60
GIACOMO BRACCHI & ANNA ALESSANDRELLO
A
B
PI. XII - Bidone pulcherrima n. gen. n. sp., MSNM Ì20624, paratype (A: x 10; B: x 16.2); Didone pulcherrima n. gen. n. sp., MSNM Ì25457, paratype (C: x 2; D: x 10.4).
xw
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRIPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
61
PI. XIII - Paleoaphrodite libanotica n. sp., MSNM il 6288, holotype (x 1 1.3).
62
GIACOMO BRACCHI & ANNA ALESSANDRELLO
PI. XIV - Ferragutia cenomaniana n. gen. n. sp., MSNM Ì8340, holotype, (A: x 60; B: x 70); Ferragiitia cenomaniana n. gen. n. sp MSNM il 2426, paratype (C: x 4; D: x 60).
PALEODIVERSITY OF THE FREE-LIVING POLYCHAETES (ANNELIDA, POLYCHAETA) AND DESCRJPTION OF NEW TAXA FROM THE UPPER CRETACEOUS OF LEBANON
63
PI. XV - Eunicites dìopatroides n. sp., MSNM Ì23084, holotype (x 4)
GIACOMO BRACCHI & ANNA ALESSANDRELLO
PI. XVI - Teruzzia pezzolii n. gen. n. sp., MSNM Ì23419 (x 1.1)
Volume XVI
[ - CARETTO P. G., 1967 - Studio morfologico con l’ausilio del metodo statistico e nuova classificazione dei Gasteropodi pliocenici attribuibili al Murex brandaris Linneo./?/?. 1-60, 1 fig., 7 tabb., 10 tavv.
II - SACCHI VIALLI G. e CANTALUPPI G., 1967-1 nuovi fossili di Goz¬ zano (Prealpi piemontesi)./?/?. 61-128, 30 figg., 8 tavv.
1 (II - PIGORINI B., 1967 - Aspetti sedimentologici del Mare Adriatico. /?/?. 129-200, 13 figg., 4 tabb. 7 tavv.
Volume XVII
jt - PINNA G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Alpe Turati (Erba, Como). Famiglie Lytoceratidae, Nannolytoceratidae, Hammatoceratidae (excl. Phymatoceratinae) Hildoceratidae (excl. Hildoceratinae e Bouleiceratinae). /?/?. 1-70, 2 tavv. n.t., 6 figg., 6 tavv.
I - VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi del-
l’Anisico superiore di Lenna in Va! Brembana (Bergamo). /?/?. 71-142, 5 figg-, 11 tavv.
II - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Al¬
pe Turati (Erba, Como). Generi Hildoceras, Phymatoceras, Paronice- ras e Frechiella. Conclusioni generali, pp. 143-204, 2 figg., 6 tavv.
Volume XVIII
- PINNA G., 1969 - Revisione delle ammoniti figurate da Giuseppe Me¬ neghini nelle Tavv. 1-22 della «Monographie des fossiles du calcaire rouge ammonitique» (1867-1881)./?/?. 5-22, 2 figg., 6 tavv.
I - MONTANARI L., 1969 - Aspetti geologici del Lias di Gozzano (Lago
d’Orta )./?/?. 23-92, 42 figg., 4 tavv. n.t.
II - PETRUCCI F„ BORTOLAMI G. C. & DAL PIAZ G. V., 1970 - Ri¬
cerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul suo substrato cristallino./?/?. 93-169, con carta a colori al 1:40.000. 14 figg., 4 tavv. a colori e 2 b.n.
Volume XIX
- CANTALUPPI G., 1970 - Le Hildoceratidae del Lias medio delle regio¬ ni mediterranee - Loro successione e modificazioni nel tempo. Riflessi biostratigrafici e sistematici, pp. 5-46, 2 tabb. n.t.
I - PINNA G. & LEVI-SETTI F., 1971 - 1 Dactylioceratidae della Provincia
Mediterranea (Cephalopoda Ammonoidea). pp. 47-136, 21 figg., 12 tavv.
II - PELOSIO G., 1973 - Le ammoniti del Trias medio di Askìepieion (Ar-
golide, Grecia) - I. Fauna del «calcare a Ptychites» (Anisico sup.). pp. 137-168, 3 figg., 9 tavv.
Volume XX
- CORNAGGIA CASTIGLIONI O., 1971 - La cultura di Remedello. Problematica ed ergologia di una facies dell’Eneolitico Padano, pp. 5-80, 2 figg., 20 tavv.
I - PETRUCCI F., 1972 - Il bacino del Torrente Cinghio (Prov. Parma). Studio sulla stabilità dei versanti e conservazione del suolo./?/?. 81-127, 37 figg., 6 carte tematiche.
,11 - CERETTI E. & POLUZZI A., 1973 - Briozoi della biocalcarenite del Fosso di S. Spirito (Chieti, Abruzzi), pp. 129-169, 18 figg., 2 taw.
Volume XXI
- PINNA G., 1974 - 1 crostacei della fauna triassica di Cene in Val Seriana (Bergamo), pp. 5-34, 16 figg., 16 tavv.
I - POLUZZI A., 1975 - 1 Briozoi Cheilostomi del Pliocene della Val d’ Ar¬
da (Piacenza, Italia)./?/?. 35-78, 6 figg., 5 taw.
II - BRAMBILLA G., 1976 - I Molluschi pliocenici di Villalvemia (Ales¬
sandria). I. Lamellibranchi. pp. 79-128, 4 figg., 10 tavv.
Volume XXII
- CORNAGGIA CASTIGLIONI O. & CALEGARI G., 1978 - Corpus delle pintaderas preistoriche italiane. Problematica, schede, iconogra¬ fia./?/?. 5-30, 6 figg., 13 tavv.
I - PINNA G., 1979 - Osteologia dello scheletro di Kritosaurns notabilis
(Lambe, 1914) del Museo Civico di Storia Naturale di Milano (Ornithi- schia Hadrosauridae). pp. 31-56, 3 figg-, 9 tavv.
II - BIANCOTTI A., 1981 - Geomorfologia dell’Alta Langa (Piemonte
meridionale), pp. 57-104, 28 figg., 12 tabb., 1 carta ft.
Volume XXIII
- GIACOBINI G., CALEGARI G. & PINNA G., 1982 -1 resti umani fossili della zona di Arena Po (Pavia). Descrizione e problematica di una serie di reperti di probabile età paleolitica, pp. 5-44, 4 figg., 16 taw.
I - POLUZZI A., 1982 - I Radiolari quaternari di un ambiente idrotermale
del Mar Tirreno, pp. 45-72, 3 figg., 1 tab., 13 tavv.
II - ROSSI F., 1984 - Ammoniti del Kimmeridgiano superiore-Berriasiano
inferiore del Passo del Furio (Appennino Umbro-Marchigiano)./?/?. 73- 138, 9 figg., 2 tabb., 8 taw.
Volume XXIV
I - PINNA G., 1 984 - Osteologia di Drepanosaurus unguicaudatas, lepido-
sauro triassico del sottordine Lacertilia. pp. 5-28, 12 figg., 2 taw.
II - NOSOTTI S. e PINNA G., 1989 - Storia delle ricerche e degli studi sui
rettili Placodonti. Parte prima 1830-1902./?/?. 29-86, 24 figg., 12 tavv.
Volume XXV
I - CALEGARI G., 1989 - Le incisioni rupestri di Taouardei (Gao, Mali). Pro¬
blematica generale e repertorio iconografico, pp. 1-14, 9 figg., 24 taw
II - PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia funzionale
e paleoecologia del rettile placodonte Psephoderma alpinnm Meyer, 1858 . pp. 15-50, 20 figg., 9 tavv.
Ili - CALDARA R., 1990 - Revisione Tassonomica delle specie paleartiche del genere Tychius Germar (Coleoptera Curculionidae). pp. 51-218, 575 figg.
Volume XXVI
I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia, Placo-
dontia). pp. 1-21, 23 figg.
II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara preisto¬
rico: dati e interpretazioni./?/?. 25-556, 647 figg.
Ili - ANDRI E. e ROSSI F., 1993 - Genesi ed evoluzione di frangenti, cinture, barriere ed atolli. Dalle stromatoliti alle comunità di scogliera moderne./?/?. 559-610, 49 figg., 1 tav.
Volume XXVII
I - PINNA G. and GHISELIN M. edited by, 1996 - Biology as History. N.
1. Systematic Biology as an Historical Science./?/?. 1-133, 68 figs.
II - LEONARDI C. e SASSI D. a cura di, 1997 - Studi geobotanici ed en-
tomofaunistici nel Parco Regionale del Monte Barro, pp. 135-266, 122 figg., 23 tabb.
Volume XXVIII
I - BANFI E. & GALASSO G., 1998 - La flora spontanea della città di Milano alle soglie del terzo millennio e i suoi cambiamenti a partire dal 1700./?/?. 267-388, 71 figg., 30 tabb.
Volume XXIX
I - CALEGARI G., 1999 - L’arte rupestre dell’Eritrea. Repertorio ragiona¬ to ed esegesi iconografica./?/?. 1-174, 268 figg.
Volume XXX
I - PEZZOTTA F. edited by, 2000 - Mineralogy and petrology of shallow
depth pegmatites. Paper from thè First International Workshop./?/?. 1-117, 30 figs., 19 tabs.
II - PARISI B., FRANCHINO A. & BERTI A. con la collaborazione di
POTENZA B. & RUBINI D., 2000 - La Società Italiana di Scienze Naturali 1855 - 2000. Percorsi storici./?/?. 1-163, 199 figg.
Ili - DE ANGELI A. & GARASSINO A., 2002 - Galatheid, chirostylid and porcellanid decapods (Crustacea, Decapoda, Anomura) from thè Eocene and Oligocene of Vicenza (N Italy). pp. 1-31, 21 figs., 9 pls.
Volume XXXI
I - NOSOTTI S. & RIEPPEL O., 2002 - The braincase of Placodus Agas-
siz, 1833 (Reptilia, Placodontia). pp. 1-18, 15 figs.
II - MARTORELLI G., 2002 - Monografia illustrata degli uccelli di rapina
in Italia. (1895). Riedizione a cura di Fausto Barbagli./?/?. [XX] 1-216, [14] 46 figg., 4 tavv.
Ili - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphargis dalsassoi n. gen. n. sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano (Lombardy, N Italy)./?/?. 1-33, 19 figs., 1 tab., 3 pls.
Volume XXXII
I - ALESSANDRELLO A., BRACCHI G. & RIOU B., 2004 - Polychaete,
sipunculan and enteropneust worms from thè Lower Callovian (Middle Jurassic) of La Voulte-sur-Rhòne (Ardèche, France)./?/?. 1-16, 9 figs., 1 pi.
II - RIEPPEL O. & HEAD J. J., 2004 - New specimens of thè fossil snake
genus Eupodophis Rage & Escuillié, from Cenomanian (Late Creta- ceous) of Lebanon./?/?. 1-26, 13 figs., 1 tab.
Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali, Museo Civico di Storia Naturale, Corso Venezia 55 - 20121 Milano Pubblicazione disponibile al cambio