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

© 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.

Volume XVIII

I - PINNA G- 1969 - Revisione delle ammoniti figurate da Giuseppe

Meneghini nelle Tavv. 1-22 della « Monographie desfossiles dii calcai-

re muge ammonitique » (1867-1881 ). pp. 5-22. 2 Jìgg.. 6 tavv.

II - MONTANARI L„ 1969 - Aspetti geologici del Lias di Gozzano (Lago

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.

Volume XIX

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

Volume XX

I - CORNEGGIA CASTIGLIONI O., 1971 - La cultura di Remedello.

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.

Ili - CERETTI E & POLUZZI A., 1973 - Briozoi della biocalcaremte del

Fosso di S. Spirito (Chieti. Abruzzi), pp. 129-169. 18 figg.. 2 tavv.

Volume XXI

I - PINNA G.. 1974 - 1 crostacei della fauna triassica di Cene in Val Seriana

(Bergamo)./?/?. 5-34. 16 Jìgg.. 16 tavv.

II - POLUZZI A.. 1975 - 1 Briozoi Cheilostomi del Pliocene della Val

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.

Volume XXII

- 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.

I - B1ANCOTTI A.. 1981 - Geomorfologia dell'Alta Langa (Piemonte

meridionale)./?/?. 57-104. 28 Jìgg.. 12 tabb.. 1 carta f. t.

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

- PINNA G.. 1984 - Osteologia di Drepanosaurus unguicaudatus. lepido-

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

L.

NEW SPECIMENS OF THE FOSSIL SNAKE GENUS EUPODOPHIS RAGE & ESCUILLIÉ, FROM THE CENOMANIAN (LATE CRETACEOUS) OF LEBANON

5

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.

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

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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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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

-

■

i

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