soc

HARVARD UNIVERSITY

Ernst Mayr Library

of thè Museum of

Comparative Zoology

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MEMORIE

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.. y^RP^ della Società Italiana

Volume XXXIV - Fascicolo I

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della Società Italiana

di Scienze Naturali

e del Museo Civico

di Storia Naturale di Milano

ALESSANDRO GARASSINO & GÙNTER SCHWEIGERT

THE UPPER JURASSIC SOLNHOFEN

DECAPOD CRUSTACEAN FAUNA:

REVIEW OF THE TYPES

FROM OLD DESCRIPTIONS

MILANO MAGGIO 2006

Elenco delle Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

Volume I

I - CORNALIA E., 1865 - Descrizione di una nuova specie dei genere

Felis: Felis jacobìta (Com.). 9 pp., 1 tav.

II - MAGNI-GRIFFI F„ 1865 - Di una specie d 'Hippoìais nuova per l’Ita-

lia. 6 pp., I tav.

Ili - GASTALDI B., 1 865 - Sulla riescavazione dei bacini lacustri per opera

degli antichi ghiacciai. 30 pp., 2 figg-, 2 tavv.

IV - SEGUENZA G., 1865 - Paleontologia malacologica dei terreni terziarii

del distretto di Messina. 88 pp., 8 tavv.

V - GIBELLI G., 1865 - Sugli organi riproduttori del genere Verrucaria,

16 pp., 1 tav.

VI - BEGGIATO F. S., 1865 - Antracoterio di Zovencedo e di Monteviale

nel Vicentino, 10 pp., 1 tav.

VII - COCCHI I., 1865 - Di alcuni resti umani e degli oggetti di umana

industria dei tempi preistorici raccolti in Toscana. 32 pp., 4 tavv.

VIII - TARGIONI-TOZZETTI A., 1866 - Come sia fatto l’organo che fa

lume nella lucciola volante dell’Italia centrale (Luciola italica) e come

le fibre muscolari in questo ed altri Insetti ed Artropodi. 28 pp., 2 tavv.

IX - MAGGI L„ 1865 - Intorno al genere Aeolosoma. 18 pp., 2 taw.

X - CORNALIA E., 1 865 - Sopra i caratteri microscopici offerti dalle Cantari¬

di e da altri Coleotteri facili a confondersi con esse. 40 pp., 4 tavv.

Volume II

I - ISSEL A„ 1 866 - Dei Molluschi raccolti nella provincia di Pisa, 38 pp.

II - GENTILLI A., 1 866 - Quelques considérations sur l’origine des bassins

lacustres, àpropos des sondages du Lac de Come. 12 pp., 8 taw.

Ili - 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 taw.

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.

VII - COCCHI I., 1866 - L’uomo fossile nell’Italia centrale. 82 pp., 21 figg.,

4 tavv.

VIII - GARO VAGLIO S., 1866 - Manzonia cantiana, novum Lichenum

Angiocarporum genus propositum atque descriptum. 8 pp., 1 tav.

IX - SEGUENZA G., 1867 - 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 taw.

Volume III

I - EMERY C., 1873 - Studii anatomici sulla Vipera Redii. 16 pp., 1 tav.

II - GAROVAGLIO S., 1867 - Thelopsis, Belonia, Weitenwebera et Limbo-

ria, quatuor Lichenum Angiocarporum genera recognita iconibusque

illustrata. 12 pp., 2 taw.

Ili - TARGIONI-TOZZETTI A., 1 867 - Studii sulle Cocciniglie. 88 pp., 7 taw.

IV - CLAPARÈDE E. R. e PANCERI P„ 1867 - Nota sopra un Alciopide

parassito della Cydippe densa Forsk. 8 pp. 1 taw.

V - GAROVAGLIO S., 1871 - De Pertusariis Europae mediae commenta¬

no. 40 pp., 4 taw.

Volume IV

I - D’ACHIARDI A., 1868 - Corallarj fossili del terreno nummulitico del-

l’Alpi venete. Parte 1 1. 32 pp. 8 tavv.

II - GAROVAGLIO S., 1868 - Octona Lichenum genera ve 1 adhuc con¬

troversa, ve I sedis prorsus incertae in systemate, novis descriptionibus

iconibusque accuratissimis illustrata. 18 pp., 2 tavv.

III - MARINONI C., 1868 - Le abitazioni lacustri e gli avanzi di umana

industria in Lombardia. 66 pp., 5 figg., 7 taw.

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 taw.

Volume VI

I - DE ALESSANDRI G., 1897 - La pietra da cantoni di Rosignano e di

Vignale. Studi stratigrafici e paleontologici. 104 pp., 2 taw, 1 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

1 - DL-. ALESSANDRI G.. 1910 - Studi sui pesci triasici della Lombardia.

164 pp., 9 lavv.

Volume Vili

I - RI: POSSI E., 1915 - La bassa Valle della Mera. Studi petrografici e

geologici. Parte \.pp. 1-46. 5 figg., 3 tavv.

Il - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi petrogra¬

fici e geologici. Parte IL pp. 47-186. 5 figg. 9 taw.

Ili - 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./?/?.

1-164, 7 figg. 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 uma¬

ne. (Saggio di una nuova dottrina craniologica con particolare riguardo

dei principali cranii fossili)./?/?. 165-262, 7 figg., 2 taw.

Ili - DE BEAUX O. e FESTA E., 1927 - La ricomparsa del Cinghiale nel¬

l’Italia settentrionale-occidentale, pp. 263-320, 13 figg., 7 tavv.

Volume X

I - DESIO A., 1929 - Studi geologici sulla regione dell’Albenza (Prealpi

Bergamasche)./?/?. 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 taw.

Ili - SCORTECCI G., 1941-1 recettori degli Agamidi./?/?. 209-326, 80 figg.

Volume XI

I - GUIGLIA D., 1944 - Gli Sfecidi italiani del Museo di Milano (Hymen.).

pp. 1-44, 4 figg., 5 tavv.

II-III - GIACOMINI 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)./?/?. 1-70, 4 figg. 6 tavv.

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), p/?. 141-188, 2 figg., 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, pp. 1-64, 25 figg.. 9 taw., 1 carta.

II - PINNA G., 1963 - Ammoniti del Lias superiore (Toarciano) dell’Alpe

Turati (Erba, Como). Generi Mercaticeras, Pseudomercaticeras e Bro-

dieìa. pp. 65-98, 2 figg., 4 tavv.

Ili - ZANZUCCHI G„ 1963 - Le Ammoniti del Lias superiore (Toarciano)

di Entratico in Val Cavallina (Bergamasco orientale), pp. 99-146, 2

figg., 8 tavv.

Volume XIV

I - VENZO S., 1965 - Rilevamento geologico dell’anfiteatro morenico

frontale 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 tavv.

III - DI ENI 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., 1 966 - Nuova classificazione di alcuni Briozoi plioce¬

nici, precedentemente determinati quali Idrozoi del genere Hydractinia

Van Beneden. pp. 1-88, 27 figg. 9 tavv.

II - D1ENI 1. 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.

Volume XVI

I - CARETTO P. G., 1967 - Studio morfologico con l’ausilio del metodo

statistico e nuova classificazione dei Gasteropodi pliocenici attribuibili

al Murex brandaris Linneo, pp. 1-60, 1 fig., 7 tabb., 10 taw.

II - SACCHI VIALLI G. e CANTALUPPI G., 1967 - 1 nuovi fossili di Goz¬

zano (Prealpi piemontesi), pp. 61-128, 30 figg., 8 taw.

Ili - PIGORIN1 B„ 1967 - Aspetti sedimentologici del Mare Adriatico, pp.

129-200, 13 figg., 4 tabb. 7 taw.

Volume XVII

I - PINNA G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Alpe

Turati (Erba, Como). Famiglie Lytoceratidae, Nannolytoceratidae,

Hammatoceratidae (excl. Phymatoceratinae) Hildoceratidae (excl.

Hildoceratinae e Bouleiceratinae). pp. 1-70, 2 tavv. n.t., 6 figg-, 6 taw.

II - VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi del-

l’Anisico superiore di Lenna in Val Brembana (Bergamo), pp. 71-142,

5 figg., 11 taw.

Alessandro Garassino & Giinter Schweigert

MCZ

RARY

Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia

Staatliches Museum fur Naturkunde, Stuttgart

JUN - 5 2006

ARD

' RSlTl

The Upper Jurassic Solnhofen decapod crustacean fauna:

review of thè types from old descriptions

Part I. Infraorders Astacidea, Thalassinidea, and Palinura

LIBRARY

jUN - 5 2006

nV.WERSnV

Volume XXXIV - Fascicolo I

Maggio 2006

Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

INDEX

Age and setting of localities . Pag. 5

Previous work and present state of thè

art . Pag. 6

Material . Pag. 6

Systematic Palaeontology . Pag. 6

Genus Eryma v. Meyer, 1840 . Pag. 6

Genus Palaeastacus Bell, 1850 . Pag. 9

Genus Pustulina Quenstedt, 1857 . Pag. 1 1

Genus Pseudastacus Oppel, 1861 . Pag. 1 2

Genus Stenochìrus Oppel, 1861 . Pag. 14

Genus Glyphea v. Meyer, 1835 . Pag. 15

Genus Squamosoglyphea Beurlen, 1930 . Pag. 17

Genus Mecochirus Germar, 1 827 . Pag. 1 8

Genus Etallonia Oppel, 1861 . Pag. 21

Genus Maglia Miinster, 1839 . Pag. 22

Genus Orhomalus Etallon, 1861 . Pag. 24

Genus Cycleryon Glaessner, 1 965 . Pag. 24

Genus Eryon Desmarest, 1 822 . Pag. 29

Genus Rosenfeldia Garassino, Teruzzi &

Dalla Vecchia, 1996 . Pag. 30

Genus Knebelia Van Straelen, 1 922 . Pag. 3 1

Genus Palaeopentacheles v. Knebel, 1 907 . Pag. 3 1

Genus Palinurina Miinster, 1839 . Pag. 32

Genus Cancrinos Miinster, 1839 . Pag. 34

Acknowledgements . Pag. 40

References . Pag. 41

© 2006 Società Italiana di Scienze Naturali

Museo Civico di Storia Naturale di Milano

Corso Venezia ,55 - 20121 Milano

In copertina: Cycleryon propinquus (Schlotheim, 1822). Reconstruction F. Fogliazza.

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 - Maggio 2006

ISSN 0376-2726

Alessandro Garassino & Gunter Schweigert

The Upper Jurassic Solnhofen decapod crustacean fauna:

review of thè types from old descriptions

Part I. Infraorders Astacidea, Thalassinidea, and Palinura

To Giorgio Teruzzi

"Le seul et le vrai moyen d’avancer la Science est de travailler à la

description et à l ’histoire des dijferentes choses qui en font l ’objet.

Buffon.

(Discours de la manière d ’étudier et de trai ter l ’Histoire Naturelle)

Abstract - The decapod crustacean fauna from Solnhofen and Eichstàtt has not been reviewed since thè studies by Schlotheim

(1820, 1822), Germar (1827), v. Meyer (1836), Munster (1839), Oppel (1861, 1862), Woodward (1866), and v. Knebel (1907). Since

many species were described by few or poorly preserved specimens, a review of thè originai sample, today housed in various German

museum’s collections, was necessary in order to establish their trae systematic validity.

This review is divided into two parts: infraorders Astacidea Latreille, 1803, Thalassinidea Latreille, 1831, and Palinura Latreille,

1803, in thè first and superfamily Penaeoidea Rafinesque-Schmaltz, 1815, plus infraorder Caridea Dana, 1852, in thè second (Sch¬

weigert & Garassino, in press).

The originai sample of thè infraorder Astacidea Latreille, 1803, consisting of 60 specimens (originally 515), was enriched with

specimens discovered in new quarries and housed in private collections to expand morphological descriptions. This review docu-

mented thè systematic validity of thè following species: Eryma modestiforme (Schlotheim, 1822), E. veltheimii (Munster, 1839), Pa-

laeastacus fuciformis (Schlotheim, 1822), Pustulina minuta (Schlotheim, 1822) (family Erymidae Van Straelen, 1924); Pseudastacus

pustulosus (Munster, 1839) (family Protastacidae Albrecht, 1983); Stenochirus angustus (Munster, 1839), and S. mayeri Oppel, 1862

(family Stenochiridae Beurlen, 1930); Glyphea pseudoscyllarus (Schlotheim, 1822), G. tenuis Oppel, 1861, G. verrucosa Munster,

1839, and Squamosoglyphea squamosa (Munster, 1839) (family Glypheidae Winckler, 1883); Mecochirus longimanatus (Schlotheim,

1822) (family Mecochiridae Van Straelen, 1924). At thè same time, thè comparative study among thè species belonging to same genera

resulted in thè following synonymies: Eryma leptodactylina (Germar, 1 827) = E. modestiforme , and Mecochirus bajeri Germar, 1 827 =

M. longimanatus. Moreover, we recognized that Pseudastacus muensteri Oppel, 1862, must be considered thè female of P. pustulosus,

and Mecochirus brevimanus (Munster, 1839) must be considered thè female of M. longimanatus. The generic position of Glyphea

verrucosa Munster, 1839, is stili unclear.

The originai sample of thè infraorder Thalassinidea Latreille, 1831, consists of 9 specimens (originally 1 9). This review documented

thè systematic validity of thè following species: Etallonia longimana (Munster, 1839), Maglia latimana Munster, 1839 (family Axiidae

Huxley, 1879); Orhomalus deformis (Oppel, 1862) (indeterminate family). At thè same time, thè comparative study among thè species

belonging to thè same genera resulted in thè following synonymy: Maglia robusta Oppel, 1862 = M. desmarestii (Munster, 1839).

The originai sample of thè infraorder Palinura Latreille, 1 803, consisting of 44 specimens (originally 446), was enriched with speci¬

mens discovered in new quarries and housed in private collections to expand morphological descriptions. This review documented thè

systematic validity of thè following species: Cycleryon propinquus (Schlotheim, 1822), C. orbiculatus (Munster, 1839), C. elongatus

(Munster, 1 839), Eryon arctiformis (Schlotheim, 1 822), and Knebelia bilobata (Munster, 1 839) (family Eryonidae De Haan, 1841); Pa-

laeopentacheles roettenbacheri (Munster, 1839) (family Polychelidae Wood-Mason, 1874); Palinurina longipes (Munster, 1839), and P.

tenera Oppel, 1862 (family Palinuridae Latreille, 1802); Cancrinos claviger Munster, 1839 (family Cancrinidae Beurlen, 1930). At thè

same time, thè comparative study among thè species belonging to same genera resulted in thè following synonymies: Cycleryon armatus

(v. Knebel, 1 907) = C. orbiculatus (Munster, 1839), Knebelia schuberti (v. Meyer, 1836) = C. propinquus, Palinurina pygmea Miinster,

1 839 = P. longipes, Cancrinos latipes Miinster, 1 839 = C. claviger. Moreover, Cycleryon spinimanus (Germar, 1 827) is thè female of C.

propinquus thanks to thè recent review by Schweigert (2001a), Cycleryon subrotundus (Munster, 1839) is probably a juvenile stage of C.

orbiculatus, and Eryon oppeli Woodward, 1866, must be ascribed to Rosenfeldia Garassino, Teruzzi & Dalla Vecchia, 1996.

Finally, we were not able to confirm thè systematic validity of Mecochirus dubius (Munster, 1839) as a result of thè poor state of

preservation of thè originai specimens. Maglia denticulata must be considered nomen dubium because thè originai specimen belonging

to this species is lost.

Key words: Crastacea, Decapoda, lithographic limestones. Upper Jurassic, Germany.

Zusammenfassung - Die oberjurassische ZehnfuBkrebs-Fauna von Solnhofen: Neubearbeitung der Typen aus alten Beschreibun-

gen. Teil I. Unterordnungen Astacidea, Thalassinidea und Palinura.

Die Fauna dekapoder Crastaceen aus Solnhofen und Eichstàtt ist seit den Arbeiten von Schlotheim (1820, 1822), Germar (1827),

v. Meyer (1836), Munster (1839), Oppel (1861, 1862), Woodward (1866), und v. Knebel (1907) nicht mehr revidiert worden. Da viele

Arten auf wenige oder schlecht erhaltene Stiicke begrundet sind, erschien eine Neubearbeitung der Originalstticke, die heute in ver-

schiedenen Museumssammlungen aufbewahrt wurden, unerlàsslich, um deren systematische Giiltigkeit zu priifen.

Dieses Review ist in zwei Teile gegliedert: Unterordnungen Astacidea Latreille, 1803, Thalassinidea Latreille, 1831, und Palinura

Latreille, 1803, im ersten Teil und Uberfamilie Penaeoidea Rafinesque-Schmaltz, 1815, sowie Unterordnung Caridea Dana, 1852 im

zweiten Teil (Schweigert & Garassino, in press).

4

ALESSANDRO GARASSINO & GÙNTER SCHWEIGERT

Die Original-Stichprobe der Unterordnung Astacidea Latreille, 1803, bestehend aus 60 Stiicken (urspriinglich 515), wurde

unter Einschluss von neu entdeckten Exemplaren aus Privatsammlungen untersucht, um die morphologischen Beschreibungen zu

komplettieren.Das Review resultierte in der Feststellung der taxonomischen Gultigkeit fùr folgende Arten: Eryma modestiforme

(Schlotheim, 1822), E. veltheimii (Munster, 1839), Palaeastacus fuciformis (Schlotheim, 1822), und Pustulina minuta (Schlotheim,

1822) (Familie Erymidae Van Straelen, 1924); Pseudastacus pustulosus (Munster, 1839) (Familie Protastacidae Albrecht, 1983);

Stenochirus angustus (Munster, 1839), und S. mayeri Oppel, 1862 (Familie Stenochiridae Beurlen, 1930); Glyphea pseudoscyllarus

(Schlotheim, 1822), G. tenuis Oppel, 1861, G. verrucosa Munster, 1839, und Squamosoglyphea squamosa (Munster, 1839) (Familie

Glypheidae Winckler, 1883); Mecochirus longimanatus (Schlotheim, 1822) (Family Mecochiridae Van Straelen, 1924). Gleichzeitig

ergaben sich aus der vergleichenden Studie die folgenden Synonymien: Eryma leptodactylinum (Germar, 1 827) = E. modestiforme ,

und Mecochirus bajeri Germar, 1 827 = M. longimanatus. AuBerdem erkannten wir, dass Pseudastacus muensteri Oppel, 1 862, als das

Weibchen von P. pustulosus und Mecochirus brevimanus (Munster, 1839) als das Weibchen von M. longimanatus betrachtet werden

muss. Die Gattungszugehòrigkeit von “ Glyphea^ verrucosa Munster, 1839 ist noch immer unklar.

Die Original-Stichprobe der Unterordnung Thalassinidea Latreille, 1831, besteht aus 9 Stiicken (urspriinglich 19). Das Review

ergab die taxonomische Gultigkeit der folgenden Arten: Etallonia longimana (Munster, 1839), und Magila latimana Munster, 1839

(Familie Axiidae Huxley, 1879); Orhomalus deformis (Oppel, 1862) (Familie indet.). AuBerdem ergab sich die folgende Synonymien:

Magila robusta Oppel, 1862 = M. desmarestii (Munster, 1839).

Die Original-Stichprobe der Unterordnung Palinura Latreille, 1803, bestehend aus 44 Exemplaren (urspriinglich 446), wurde zur Ver-

vollstàndigung der morphologischen Beschreibung zusammen mit neu entdeckten Stiicken aus Privatsammlungen untersucht. Dabei ergab

sich die taxonomische Gultigkeit der folgenden Arten: Cycleryon propinquus (Schlotheim, 1 822), C. orbiculatus (Mtinster, 1 839), C. elon-

gatus (Munster, 1839), Eryon arctiformis (Schlotheim, 1822), und Knebelia bilobata (Miinster, 1839) (Familie Eryonidae De Haan, 1841);

Palaeopentacheles roettenbacheri (Miinster, 1 839) (Familie Polychelidae Wood-Mason, 1 874); Palinurina longipes (Munster, 1 839), und P.

tenera Oppel, 1 862 (Familie Palinuridae Latreille, 1 802); Cancrinos claviger Munster, 1 839 (Familie Cancrinidae Beurlen, 1 930). Hierbei

ergaben sich folgende Synonymien: Cycleryon armatus (v. Knebel, 1907) = C. orbiculatus (Miinster, 1839), Knebelia schuberti (v. Meyer,

1836) = C. propinquus, Palinurina pygmea Munster, 1839 = P. longipes, Cancrinos latipes Munster, 1839 = C. claviger,. Dariiber hinaus

erkannten wir aufgrund der kiirzlich erfolgten Neubearbeitung durch Schweigert (200 la), dass Cycleryon spinimanus (Germar, 1827) das

Weibchen zu C. propinquus darstellt. Cycleryon subrotundus (Miinster, 1 839) ist vermutlich nur ein Jugendstadium von C. orbiculatus, und

Eryon oppeli Woodward, 1 866, muss zur Gattung Rosenfeldia Garassino, Teruzzi & Dalla Vecchia, 1 996, gestellt werden.

SchlieBlich waren wir aufgrund der schlechten Erhaltung der Originalstiicke nicht in der Lage, die taxonomische Gultigkeit von

Mecochirus dubius (Munster, 1 839) zu bestàtigen. Magila denticulata muss auBerdem als nomen dubium angesehen warden, da keine

Syntypen dieser Art mehr vorhanden sind, und die urspriingliche Beschreibung ungenugend ist.

Schlusselwòrter: Crustacea, Decapoda, Plattenkalke, Oberjura, Deutschland.

Riassunto - La fauna a crostacei decapodi del Giurassico superiore di Solnhofen: revisione degli esemplari tipo dalle descrizioni

originali. Parte I. Infraordini Astacidea, Thalassinidea e Palinura.

La fauna a crostacei decapodi di Solnhofen e Eichstatt non è stata più oggetto di revisione dopo gli studi condotti da Schlotheim

(1820, 1822), Germar (1827), v. Meyer (1836), Munster (1839), Oppel (1861, 1862), Woodward (1866) e v. Knebel (1907). Visto che

molte specie sono state descritte sulla base di pochi esemplari o di esemplari in cattivo stato di conservazione, una revisione del cam¬

pione originale, attualmente depositato presso alcuni musei tedeschi era quanto mai necessaria per stabilire la validità sistematica delle

specie descritte. Questa revisione ha avuto come oggetto di studio sia i decapodi natanti che quelli reptanti.

La revisione viene divisa in due parti: gli infraordini Astacidea Latreille, 1803, Thalassinidea Latreille, 1831 e Palinura Latreille,

1803, nella prima e la superfamiglia Penaeoidea Rafinesque-Schmaltz, 1815 e Tinfraordine Caridea Dana, 1852, nella seconda (Sch¬

weigert & Garassino, in stampa).

11 campione originale delTinfraordine Astacidea Latreille, 1803, formato da 60 esemplari (originariamente 515), è stato integrato

con esemplari rinvenuti in nuove località e depositati presso collezioni private così da approfondire le descrizioni morfologiche. Questa

revisione ha documentato la validità sistematica delle seguenti specie: Eryma modestiforme (Schlotheim, 1822), E. veltheimii (Munster,

1839), Palaeastacus fuciformis (Schlotheim, 1822) e Pustulina minuta (Schlotheim, 1822) (famiglia Erymidae Van Straelen, 1924);

Pseudastacus pustulosus (Munster, 1839) (famiglia Protastacidae Albrecht, 1983); Stenochirus angustus (Miinster, 1839) e S. mayeri

Oppel, 1862 (famiglia Stenochiridae Beurlen, 1930); Glyphea pseudoscyllarus (Schlotheim, 1822), G. tenuis Oppel, 1861, G. verru¬

cosa Munster, 1839 e Squamosoglyphea squamosa (Munster, 1839) (famiglia Glypheidae Winckler, 1883); Mecochirus longimanatus

(Schlotheim, 1822) (famiglia Mecochiridae Van Straelen, 1924). Lo studio comparativo tra le specie appartenenti allo stesso genere ha

messo in evidenza le seguenti sinonimie: Eryma leptodactylina (Germar, 1827) = E. modestiforme e Mecochirus bajeri Germar, 1827 =

M. longimanatus. Inoltre, riteniamo che Pseudastacus muensteri Oppel, 1862, sia la femmina di P. pustulosus e Mecochirus brevimanus

(Munster, 1839) sia la femmina di M. longimanatus. La posizione sistematica di Glyphea verrucosa Miinster, 1839, è ancora incerta.

Il campione originale delTinfraordine Thalassinidea Latreille, 1831, si compone di 9 esemplari (originariamente 19). Questa revisione

ha documentato la validità sistematica delle seguenti specie: Etallonia longimana (Miinster, 1839) e Magila latimana Miinster, 1839

(famiglia Axiidae Huxley, 1879); Orhomalus deformis (Oppel, 1862) (famiglia indeterminata). Lo studio comparativo tra le specie apparte¬

nenti allo stesso genere ha messo in evidenza la seguente sinonimia: Magila robusta Oppel, 1862 = M. desmarestii (Munster, 1839).

Il campione originale delTinfraordine Palinura Latreille, 1803, formato da 44 esemplari (originariamente 446), è stato integrato

con esemplari rinvenuti in nuove località e depositati presso collezioni private così da approfondire le descrizioni morfologiche.

Questa revisione ha documentato la validità sistematica delle seguenti specie: Cycleryon propinquus (Schlotheim, 1 822), C. orbicu¬

latus (Munster, 1839), C. elongatus (Munster, 1839), Eryon arctiformis (Schlotheim, 1822) e Knebelia bilobata (Miinster, 1839)

(famiglia Eryonidae De Haan, 1841); Palaeopentacheles roettenbacheri (Munster, 1839) (famiglia Polychelidae Wood-Mason, 1874);

Palinurina longipes (Munster, 1839) e P. tenera Oppel, 1862 (famiglia Palinuridae Latreille, 1802); Cancrinos claviger Munster,

1839 (famiglia Cancrinidae Beurlen, 1930). Lo studio comparativo tra le specie apparteneti allo stesso genere ha messo in evidenza

le seguenti sinonimie: Cycleryon armatus (v. Knebel, 1907) = C. orbiculatus (Munster, 1839), Knebelia schuberti (v. Meyer, 1836) =

C propinquus, Palinurina pygmea Munster, 1839 = P. longipes, Cancrinos latipes Miinster, 1839 = C. claviger. Inoltre, riconosciamo

che Cycleryon spinimanus (Germar, 1827) è la femmina di C. propinquus grazie alla recente revisione di Schweigert (200 la), che

Cycleryon subrotundus (Miinster, 1839) è un probabile stadio giovanile di C. orbiculatus e che Eryon oppeli Woodward, 1866, deve

essere attribuito a Rosenfeldia Garassino, Teruzzi & Dalla Vecchia, 1996.

Infine, non siamo in grado di confermare la validità sistematica di Mecochirus dubius (Munster, 1839) a causa del cattivo stato

di conservazione degli esemplari originali. Magila denticulata deve essere considerata nomen dubium in quanto l’unico esemplare

originale appartenente a questa specie è andato perduto.

Parole chiave: Crustacea, Decapoda, calcare litografico, Giurassico superiore, Germania.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

5

AGE AND SETTING OF LOCALITIES

The lithographic limestones of thè Upper Jurassic

in southern Germany comprise a large set of localities

of different ages and settings spanning an area of sev-

eral hundreds of kilometres (Fig. 1). The localities in

southern Franconia are often summarised as “Solnhofen

Lithographic Limestones”. Many fossils both in old col-

lections and new material from fossil traders are labelled

as coming from “Solnhofen”. This is very misleading and

makes difficult to recognise thè evolutionary trends and

locai differences in palaeoecology. Only a very intimate

knowledge of thè locai lithologies permits distinguishing

thè exact origin of thè material. Recently, however, strati-

graphically well-placed material comes from several

scientific excavations. Besides, there exist private collec-

tions which focus on special sites. The biostratigraphic

framework of thè late Kimmeridgian (Upper Jurassic)

was worked out by one of thè authors (Schweigert et al,

1 996); however knowledge of thè succession of ammo¬

nite faunas from thè Tithonian is stili in a preliminary

stage.

The oldest lithographic limestones of southern Ger¬

many, most recently discovered and dated by ammonites,

occur in northem Franconia, at Wattendorf near Bamberg.

They are of late Kimmeridgian age (Pseudomutabilis

Zone). Another decapod-bearing locality, not yet pre-

cisely dated, occurs in thè surroundings of Regensburg

(Ebenwies). A more important fossil site nearby is that of

Brunn, a papery shale of late Kimmeridgian age deposited

in a very shallow lagoon surrounded by calcareous green

algae meadows, coral reefs, and small islands. Somewhat

younger, but stili of late Kimmeridgian age is thè Nus-

plingen Lithographic Limestone in SW Germany, which

is famous for its richness in fossil sharks and decapod

crustaceans. This limestone was deposited in about 100

m deep lagoon surrounded by sponge-microbial mounds

and islands. In thè east of Eichstatt in Franconia, several

localities yielded decapod crustaceans from lithographic

limestones around thè Kimmeridgian/Tithonian boundary,

depending on thè definition of thè lower boundary of thè

Tithonian. Excavations by thè Jura-Museum (Eichstatt)

in thè siliceous lithographic limestones of Zandt and thè

more biodetritic limestones of thè nearby Òchselberg yield

a very diverse decapod fauna in an excellent state of pres-

ervation, among them several new taxa. Other decapod

faunas similar in age to those of Zandt come from quarries

in thè north of Eichstatt (e.g. Blumenberg, Schemfeld,

Wintershof). Many old taxa were first described from

this area. The “classical” lithographic limestones of Sol¬

nhofen and Langenaltheim have an early Tithonian age,

clearly younger than those of thè nearby Eichstatt district.

Apart from mass occurrences of planktonic crinoids ( Sac -

cocoma ), these limestones are very poor in fossils. The

vertebrate fauna and other criteria suggest a more pelagic

setting. Both thè sections of thè Eichstatt and Solnhofen

districts of micritic limestones deposited in larger shallow

basins, thè margins of which are poorly known, because

today for economie reasons only thè purest limestones in

thè centrai parts of thè basins are quarried. The Solnhofen

Lithographic Limestone Formation is usually overlain by

a hardground or another disconformity followed by thè

more biodetrital and silica-rich Moemsheim Formation

which also has yielded many decapod crustaceans.

Fig. 1 - Geographic map of thè fossiliferous localities.

6

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

PREVIOUS WORKS AND PRESENT STATE OF THE ART

The bulk of thè decapod fauna of thè lithographic

limestones in southern Germany is described in mono-

graphs or short papers of thè 19th century (Schlotheim,

1820, 1822; Germar, 1827; v. Meyer, 1836; Miinster,

1839; Oppel, 1861, 1862; Woodward, 1866). Around thè

millennium, thè previous discoveries of modem eryonids

during deep sea expeditions encouraged thè study of fossil

forms (Peiser, 1904; v. Knebel, 1907). In thè first half of

thè 20th century, both Beurlen and Glaessner, thè lead-

ing authorities of fossil crustaceans in Europe, almost

excluded thè decapods from thè lithographic limestones

from their scientific works. Recently, about 56 valid taxa

from Franconia and four others from thè isolated local-

ity of Nusplingen in Swabia are described. Only a few of

them were briefly revised by Forster (1966, 1967, 1973,

1977). After his sudden death in 1987, thè study of fossil

decapods from Germany declined. From thè late 1990

onwards, both scientific excavations in eastem Bavaria

(Brunn), southern Franconia (Schamhaupten) and in Nus¬

plingen (Swabia) re-stimulated thè research activities of

decapods (Garassino & Schweigert, 2004; Garassino et

al., 2005; Polz, 1999, 2000; Schweigert 200 la, 200 lb,

200 le, 2002, 2003, 2004a, 2004b; Schweigert & Dietl,

1999; Schweigert & Frattigiani, 2005; Schweigert & Ga¬

rassino, 2003, 2004, 2005a, 2005b, in press; Schweigert &

Ròper, 2001; Schweigert et al., 2000, 2003). At that time it

became obvious that both for palaeoenvironmental recon-

struction and systematic palaeontology, it was necessary to

separate thè fossil material of each locality. Another strong

pulse was given by thè monographic documentation of

“Solnhofen” fossils which were stored in various muse-

ums around thè globe and in several excellent private col-

lections (Frickhinger, 1994, 1999). Frickhinger’s aim was

a complete documentation of all taxa of animals and plants

occurring in thè “Solnhofen” limestones. From this other-

wise unavailable data pool thè existence of many new taxa

became clear. Additionally, good material of rare or poorly

preserved taxa carne to light often allowing a review.

MATERIAL

This review is based on 113 specimens among thè

infraorders Astacidea Latreille, 1803, Thalassinidea

Latreille, 1831, and Palinura Latreille, 1803. The most

part of thè originai sample, 980 specimens, was destroyed

during World War II. The survived specimens, today

housed in thè Museum fur Naturkunde der Humboldt-

Universitàt in Berlin, thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich, thè Geological

Institute and Museum of thè University of Halle, and

Natur-Museum Senckenberg in Frankfurt am Main, were

reviewed in order to confimi their systematic placement.

Since it was impossible to clean thè originai specimens of

some species in order to describe some morphological charac-

ters not reported in thè originai descriptions, some specimens

from Frattigiani, Rudel, Tischlinger, Wulf, and Koschny

private collections were used to expand thè morphological

descriptions of thè following species: E. modestiforme, P.

minuta, P. muensteri, G. tenuis, S. angustus, C. elongatus, P.

roettenbacheri, P. longipes, P tenera, and M. latimana.

One specimen in lateral view, housed in thè Bayer¬

ische Staatssammlung fur Palàontologie und Geologie in

Munich, was used to describe for thè first time thè shape

of abdominal somites and thè tail fan of Cancrinos clav-

iger.

Finally, thè total length of thè specimens is taken dor-

sally from thè tip of thè rostmm to thè distai extremity

of thè telson and thè three dimensionai reconstructions of

thè studied species reported thè morphological characters

observed in thè originai and new specimens belonging to

thè studied species.

The systematic palaeontology used in this paper follows

thè recent classification proposed by Martin & Davis (2001 ).

Acronyms. MNHB: Museum fur Naturkunde der

Humboldt-Universitàt (Berlin, Germany); BSPG: Bay¬

erische Staatssammlung fur Palàontologie und Geologie

(Munich, Germany); SMF: Natur-Museum Senckenberg

(Frankfurt am Main, Germany); SMNS: Staatliches

Museum fur Naturkunde (Stuttgart, Germany); IGH:

Geological Institute and Museum of thè University

(Halle, Germany); BM: British Naturai History Museum

(London, Great Britain).

SYSTEMATIC PALAEONTOLOGY

Infraorder Astacidea Latreille, 1803

Family Erymidae Van Straelen, 1924

Genus Eryma v. Meyer, 1 840

Type species: Macrourites modestiformis Schlotheim,

1822

Solnhofen species: Eryma modestiforme (Schlotheim,

1822); Eryma veltheimii (Miinster, 1839)

Eryma modestiforme (Schlotheim, 1 822)

Fig. 8, PI. I (fig. 1), PI. XI (fìgs. 1,2)

1822 - Macrourites modestiformis Schlotheim; p. 29, PI. 2 (fig. 3)

1822 - Macrourites modestiformis Schlotheim in Desmarest; PI. 1 1 (fig. 5)

1827 - Astacus leptodaetylus Germar; p. 100, PI. 1 (fig. 4), nov. syn.

1839 - Glyphea modestiformis Miinster; p. 19, PI. 9 (figs. 1-3)

1839 - Glyphea crassula Miinster; p. 17, PI. 8 (fig. 5)

1839 - Glyphea elongata Miinster; p. 18, PI. 8 (fig. 8) (not figs. 11-12)

1 839 - Glyphea laevigata Miinster; p. 20, PI. 9 (figs. 5-7)

not 1839 - Glyphea Veltheimii Miinster; p. 22, PI. 10 (fig. 1) = Eryma

veltheimii, holotype

1852 - Astacus modestiformis Quenstedt; p. 268, PI. 20 (fig. 15)

1855 - Glyphea modestiformis Miinster in Fraas; p. 94

1857 - Glyphea modestiformis Miinster in Quenstedt; p. 807, PI. 99

(fig. 29)

not 1857 - Glyphea Veltheimii Miinster in Quenstedt; p. 807, PI. 100

(fig. 1 ) = Eryma westphali Schweigert, Dietl & Ròper, 2000

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

7

1861 - Eryma modestiformis (Schlotheim) in Oppel; p. 358

1861 - Ery’ma leptodactylina (Germar) in Oppel; p. 358

not 1861 - Eryma Veltheimi Miinster in Oppel; p. 358

not 1861 - Eryma major Oppel; p. 358 = Erymastacus major , from

Nusplingen

not 1861 - Eryma punctata Oppel; p. 359 = Ery’ma punctatum, from

Nusplingen

1862 -Eryma modestiformis (Schlotheim) in Oppel; p. 33, PI. 6 (figs. 5-8)

1 862 - Eryma leptodactylina (Germar) in Oppel; p. 35, PI. 7 (figs. 1-4)

not 1862 - Ery’ma Veltheimi Miinster in Oppel; p. 36, PI. 7 (fig. 5) = E.

veltheimii

not 1 862 - Eryma major Oppel in Oppel; p. 37, PI. 8 (fig. 3) = Eryma¬

stacus major

not 1862 - Eryma punctata Oppel in Oppel; p. 38, PI. 8 (fig. 4) = E.

punctatum

1867 - Astacus modestiformis Quenstedt; p. 320, PI. 25 (fig. 15)

1885 - Astacus modestiformis Quenstedt; p. 409, PI. 32 (fig. 6)

1904 - Eryma modestiformis (Schlotheim) in Walther; p. 174

1904 - Ery’ma leptodactylina (Germar) in Walther; p. 174

not 1904 - Eryma Veltheimi Miinster in Walther; p. 174 = E. westphali

Schweigert, Dietl & Ròper, 2000

not 1908 - Eryma veltheimi Munster in Engel; p. 470 = E. westphali

Schweigert, Dietl & Ròper, 2000

1925 - Eryma modestiformis (Schlotheim) in Van Straelen; p. 272

1925 - Eryma leptodactylina (Germar) in Van Straelen; p. 273

1928 - Eryma modestiformis (Schlotheim) in Beurlen; p. 164

1 928 - Eryma pseudoventrosa Beurlen; p. 163

not 1928 - Eryma punctata Oppel in Beurlen; p. 164 = E. punctatum

1928 - Clytia leptodactylina (Germar) in Beurlen; p. 170

not 1928 - Clytia veltheimi (Miinster) in Beurlen; p. 170 = E. veltheimii

1929 - Clytia leptodactylus (Germar) in Glaessner; p. 116

1929 - Eryma modestiformis (Schlotheim) in Glaessner; p. 156

1929 - Eryma pseudoventrosa Beurlen in Glaessner; p. 158

1931 - Eryma modestiformis (Schlotheim) in Roll; p. 75

1961 - Clytia leptodactylus (Germar) in Kuhn; p. 22

1961 - Eryma modestiformis (Schlotheim) in Kuhn; p. 22

1965 - Eryma modestiformis (Schlotheim) in Forster; p. 138, PI. 2 (fig. 5)

1966 - Eryma modestiformis (Schlotheim) in Forster; p. 118, PI. 16

(figs. 8?-9), PI. 17 (fig. 1)

1969 - Eryma modestiformis (Schlotheim) in Malz; p. 298, Text-fig. 7

1969 - Eryma modestiformis (Schlotheim) in Glaessner; R 455, Text-

fig. 258/4

1 994 - Eryma modestiformis (Schlotheim) in Frickhinger; p. 118, Text-

figs. 201-202

1 996 - Eryma in Ròper, Rothgaenger & Rothgaenger; Text-fig. 90

1997 - Eryma elongata (Munster) in Dietl, Dietl, Kapitzke, Rieter,

Schweigert, Ilg & Hugger; PI. 2 (fig. 2)

1998 - Eryma modestiformis (Schlotheim) in Ròper & Rothgaenger; p.

50, Text-fig. 74, PI. 1 (fig. 86)

2000 - Eryma modestiformis (Schlotheim) in Ròper, Rothgaenger &

Rothgaenger; p. 91, Fig. 108

2000 - Eryma modestiforme (Schlotheim) in Schweigert, Dietl &

Ròper; p. 4, Text-fig. la, PI. 1 (figs. 1-5)

2001 - Eryma modestiforme (Schlotheim) in Dietl & Schweigert; p.

71, Text-fig. 86

2005b - Eryma modestiforme (Schlotheim) in Schweigert & Garassino;

p. 498

Diagnosis: carapace cylindrical laterally flattened; cer-

vical groove deep and moderately inclined, not strongly

sinuous and lying almost in thè middle part of carapace;

postcervical and branchiocardiac grooves nearly parallel

and not joined; gastro-orbital groove weak; antennal and

ventral grooves deep; hepatic notch and muscle adductor

testis deep; rostrum short with three small supralateral

teeth; antennal spine well developed; pereiopods I-III

chelate; pereiopod I larger and stronger than pereiopods

II-III; uropodal exopod with diaeresis.

Holotype: Macrourites modestiformis (= Eryma

modestiforme ), MNHB K 37 MB (Schlotheim, 1822 - PI.

2, fig. 3; length of thè specimen: 2.5 cm), housed in thè

Museum fur Naturkunde der Humboldt-Universitàt in

Berlin.

Our studies revealed that Astacus leptodactylus from

Solnhofen and figured by Germar (1827 - PI. 1, fig. 4) is

synonym with Eryma modestiforme , as discussed below.

This specimen, thè holotype of Germar’s species, was

said to be lost (Forster, 1966), but recently reappeared in

thè collection of thè Geological Institute and Museum of

thè University of Halle (comm. by N. Hauschke, Halle).

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - lower Tithonian).

Type locality: Eichstàtt.

Material: Oppel (1862) reported 80 specimens

belonging to this species, so divided: 27 from thè Reden-

bacher collection and 53 from thè collection of thè Bayer-

ische Staatssammlung fur Palàontologie und Geologie in

Munich. We studied of this originai sample thè holotype

by Schlotheim, housed in thè Museum fur Naturkunde der

Humboldt-Universitàt in Berlin, and 1 1 additional speci¬

mens, housed in thè Bayerische Staatssammlung fur Palà¬

ontologie und Geologie in Munich in which we identified

seven specimens, illustrated by Munster (1839) and two

specimens, illustrated by Oppel (1862). Two specimens

(BSPG AS Vili 80, AS I 623) were not illustrated by thè

authors. We report thè list of thè specimens, studied by

Munster and Oppel:

BSPG AS VII 191 - Munster (1839), PI. 8 (fig. 12)

- length of thè specimen: 1 .5 cm

BSPG AS VII 193 - Munster (1839), PI. 8 (fig. 5)

- length of thè specimen: 3 cm

BSPG AS VII 194 - Munster (1839), PI. 9 (fig. 5)

- length of thè specimen: 4 cm

BSPG AS VII 195 - Munster (1839), PI. 9 (fig. 2)

- length of thè specimen: 3 cm

BSPG AS VII 196 - Munster (1839), PI. 9 (fig. 3)

- length of thè specimen: 2 cm

BSPG AS VII 197 - Munster (1839), PI. 9 (fig. 6)

- length of thè specimen: 2 cm

BSPG AS VII 198 - Munster (1839), PI. 9 (fig. 7)

- length of thè specimen: 2 cm

BSPG AS Vili 81 - Oppel (1862), PI. 6 (fig. 6)

- incomplete specimen

BSPG AS Vili 82 - Oppel (1862), PI. 6 (fig. 7)

- length of thè specimen: 2 cm

Oppel (1862) reported 61 specimens which he

assigned to Eryma leptodactylina (Germar, 1827), so

divided: 18 from thè Redenbacher collection and 43

from thè collection of thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich. Today, we know

of this originai sample eight specimens, housed in Bay¬

erische Staatssammlung fur Palàontologie und Geologie

in Munich, in which we identified four specimens, illus¬

trated by Munster (1839) and four specimens, illustrated

by Oppel (1862). We report thè list of thè specimens,

studied by Munster and Oppel:

BSPG AS VII 187 - Munster (1839), PI. 8 (fig. 9)

- length of thè specimen: 4 cm

8

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

BSPG AS VII 188 - Munster (1839), PI. 8 (fig. 8)

- length of thè specimen: 3 cm

BSPG AS VII 189 - Munster (1839), PI. 9 (fig. 1)

- length of thè specimen: 4 cm

BSPG AS VII 190 - Munster (1839), PI. 8 (fig. 10)

- length of thè specimen: 4 cm

BSPG AS VI 15 - Oppel (1862), PI. 7 (fig. 3)

- length of thè specimen: 6 cm

BSPG AS VI 16 - Oppel (1862), PI. 7 (fig. 4)

- length of thè specimen: 8 cm

BSPG AS VI 17 - Oppel (1862), PI. 7 (fig. 2)

- length of thè specimen: 4 cm

BSPG AS VIII 79 - Oppel (1862), PI. 7 (fig. 1)

- length of thè specimen: 3 cm

One specimen (n. cat. 9333 - Wulf collection) was

used to describe thè tail fan of this species, not preserved

in thè originai sample.

Description. Small-sized erymid with strongly tuber-

culate exoskeleton.

Carapace. Carapace cylindrical, laterally flattened.

Cervical groove deep, lying almost at thè middle part of

thè carapace. Rostrum short with three small supralateral

teeth directed forward. Postcervical and branchiocardiac

grooves nearly parallel and not joined near their lower

ends. Gastro-orbital groove deep. Antennal and ventral

grooves deep. Hepatic notch and muscle adductor testis

deep. Antennal spine well developed. Ocular incision

large and deep. Spines short and strong located in gastric

region. Pits small located in antennal, cardiac, branchial,

and pterygostomial regions.

Abdomen. Somites I-V subrectangular of equal length

with small pits on tergal surface. Somite VI subsquare.

Pleurae of somites I-VI with pointed distai extremity.

Telson subtriangular with rounded distai extremity, deep

longitudinal median groove, a pair of small median spines

and a pair of small spines located in thè distai part of lateral

margins. Protopod subrectangular. Uropodal endopod with

a strong median longitudinal carina and a small spine in thè

distai part of outer margin. Uropodal exopod with a strong

median longitudinal carina, two distai spines on thè outer

margin (one longer and one shorter) and rounded diaeresis

with serrate upper margin having one longer median spine.

Cephalic appendages. Eyestalk short. 3rd maxilliped

elongate having thè first proximal article with two strong

distai spines and second proximal article with one strong

distai spine, all located in thè lower margin. Antennular

peduncles thin of which thè distai one carries two elon¬

gate multiarticulate flagella of equal length. Antennal

articles strong and stout of which thè distai one carries a

strong elongate multiarticulate flagellum, twice thè body

length. Scaphocerite elongate and triangular with basicer-

ite having a strong distai spine in thè lower margin.

Thoracic appendages. Pereiopod I with short and stout

chela (ratio propodus/fixed finger = 1/1). Movable and

fixed fingers of equal length with curved distai extremity.

Inner margins of movable and fixed fingers with small

rounded teeth. Surface of merus, carpus, propodus and

movable finger covered with small pits and tubercles.

Chela of pereiopod III longer than chela of pereiopod

II. Pereiopods IV-V achelate. Pereiopod IV longer and

stronger than pereiopod V. Lower margin of propodus of

pereiopod IV with a row of downcurved spines.

Abdominal appendages. Pleopods with an unsegmented

peduncle which carries two elongate multiarticulate flagella.

Discussion. Forster (1966) reviewed this genus,

describing 35 species and pointing out that E. mode-

stiforme (Schlotheim, 1822) differs from thè other spe¬

cies of thè same genus by its typical midiine position

of thè cervical groove. Among thè species described by

Forster (1966), E. verrucosa (Munster, 1839) cannot be

assigned to a genus because of thè incompleteness of thè

lectotype (see Pustulina minuta in this paper). Moreover,

Forster (1966) did not include among thè studied species

of Eryma, thè species from thè Upper Cretaceous of thè

United States, described by Rathbun (1923, 1926, 1935):

E. americana Rathbun, 1923 (Black Creek Fm., N Caro¬

lina), E. flecta Rathbun, 1926 (Ripley Fm., Tennessee)

and E. stantoni Rathbun, 1935 (Ripley Fm., Alabama);

thè species from thè Upper Cretaceous (Cenomanian) of

Lebanon, described by Roger (1946): E. cretacea Roger,

1946; thè species from thè Upper Jurassic (Kimmeridgian

- Tithonian) of Madagascar described by Secretan ( 1 964):

E. madagascariensis Secretan, 1964, E. granulifera

Secretan, 1964, and E1, australis (Secretan, 1964).

Feldmann (1979), Garassino (1996) and Schweigert

et al. (2000) described three new species to add to

those described by thè above-mentioned authors: E.

foersteri Feldmann, 1979 (United States - Callovian),

E. meyeri Garassino, 1996 (Italy - Sinemurian), and E.

westphali Schweigert, Dietl & Ròper, 2000 (Germany

- Kimmeridgian). Finally, Crónier & Courville (2004)

described E. burgundiaca (France - Callovian), and

Feldmann & Titus (2006) described E. jungostrix (United

States - Oxfordian).

Today, Eryma v. Meyer, 1 840, known from thè Lower

Jurassic (Sinemurian) to thè Upper Cretaceous (Cenoma¬

nian) of Europe (Germany, France, England, Russia and

Italy), Africa (Morocco), Iran, Madagascar, Lebanon,

Antarctic Peninsula, Canada, United States, and South

America (Argentina), includes 47 species.

Oppel (1862), describing E. leptodactylina (Germar,

1827), pointed out that this species was similar to E.

modestiforme (Schlotheim, 1822) and E. veltheimii

(Munster, 1839), but it was different in exhibiting larger

chelae on pereiopod I. However, thè study of thè eight

originai specimens by Munster and Oppel, and thè very

poorly preserved holotype of Germar’s species allowed

recognition of some morphological characters, such

as thè midiine position of thè cervical groove, rostrum

with three small supralateral teeth, short and stout chela

of pereiopod I not as long as described by Oppel (ratio

propodus/fixed finger =1/1, apart from thè onthogenetic

stage of thè studied specimens) and telson with a deep

median longitudinal groove and a small spine in thè

median part of thè lateral margin. Thus, it may be useful

to include E. leptodactylina in E. modestiforme. These

characters distinguish E. modestiforme from E. veltheimii

having a rostrum with three strong supralateral teeth and

very elongate chela of pereiopod I (ratio propodus/fixed

finger = 2/1) (see E. veltheimii in this paper).

Eryma veltheimii (Munster, 1839)

Fig. 8, PI. I (fig. 2), PI. XI (figs. 3, 4)

1839 - Glyphea Veltheimii Munster; p. 22, PI. 10 (fig. 1)

1 848 - Eryma Veltheimi (Munster) in Bronn; p. 468

1 853 - Glyphea Veltheimi Munster in Frischmann; p. 28

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

9

not 1855 - Glyphea Veltheimi Mùnster in Fraas; p. 94 = Eryma west-

phali Schweigert, Dietl & Ròper, 2000

not 1857 - Glyphea Veltheimii Miinster in Quenstedt; p. 807, PI. 100

(fig. 1 ) = £. westphali Schweigert, Dietl & Ròper, 2000

1861 - Eryma Veltheimi (Miinster) in Oppel; p. 358

1862 - Eryma Veltheimi (Miinster) in Oppel; p. 36, PI. 7 (fig. 5)

1996 - Eryma in Ròper, Rothgaenger & Rothgaenger; Text-fig. 91

2000 - Eryma veltheimi (Miinster) in Schweigert, Dietl & Ròper; p. 7,

Text-fig. le, PI. 4 (figs. 1-3)

2005b - Eryma veltheimii (Miinster) in Schweigert & Garassino; p. 498

Diagnosis; carapace cylindrical laterally flattened; cer-

vical groove deep, and moderately inclined not strongly

sinuous; postcervical and branchiocardiac grooves nearly

parallel and joined near their lower ends; gastro-orbital

groove weak; antennal and ventral grooves deep; hepatic

notch and muscle adductor testis deep; rostrum short

with three strong supralateral teeth; antennal spine well

developed; pereiopods I-III chelate; pereiopod I larger

and stronger than pereiopods II-III; uropodal exopod with

diaeresis.

Holotype by monotypy: BSPG AS VII 186 a-b (Mun-

ster, 1839 - PI. 10, fig. 1; Oppel, 1862 - PI. 7, fig. 5),

housed in thè Bayerische Staatssammlung fìir Palàontolo-

gie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - lower Tithonian).

Type locality: labelled as coming from Eichstàtt;

however, based upon thè lithology of thè rock matrix, thè

specimen comes from thè Kapfelberg quarry near Kel-

heim (Schweigert et al., 2000).

Material: Oppel (1862) reported one specimen,

ascribed to this species, and housed in thè Bayerische

Staatssammlung fur Palàontologie und Geologie in

Munich where, today, it is stili preserved.

BSPG AS VII 186 a-b -Miinster (1839), PI. 10 (fig. 1)

- length of thè specimen: 8 cm (thè specimen is preserved

as part and counter-part; Miinster figured thè specimen

with thè tail fan opens in dorsal view; in realty thè tail

fan preserves thè right uropods overlapped onto thè last

abdominal somites, as figured by Oppel).

BSPG AS VII 186 a-b - Oppel (1862), PI. 7 (fig. 5)

Description. Large erymid with strongly tuberculate

exoskeleton.

Carapace. Carapace cylindrical laterally flattened.

Cervical groove deep in thè anterior part of thè carapace.

Rostrum short with three strong supralateral teeth directed

forward. Postcervical and branchiocardiac grooves nearly

parallel and joined near their lower ends. Gastro-orbital

groove weak. Antennal and ventral grooves deep. Hepatic

notch and muscle adductor testis deep. Antennal spine

well developed. Ocular incision large and deep. Spines

short and strong located in gastric region. Pits small

located in antennal, cardiac, branchial, and pterygosto-

mial regions.

Abdomen. Somites I-V subrectangular of equal length

with small pits on tergal surface. Somite VI subsquare.

Pleurae of somites I-V with pointed distai extremity.

Telson triangular with rounded distai extremity and

deep median longitudinal groove. Protopod subrectan¬

gular. Uropodal endopod with a strong median longitu¬

dinal carina. Uropodal exopod with a strong longitudinal

median carina, one small distai spine on thè outer margin,

and rounded diaeresis.

Cephalic appendages. Eyestalk short. 3rd maxilliped,

antennulae and antennae not preserved.

Thoracic appendages. Pereiopod I with elongate chela

(ratio propodus/fixed finger = 2/1). Movable and fixed

fingers of equal length with curved distai extremity, and

edentate inner margins. Upper margin of propodus with

one strong distai spine. Surface of merus, carpus, pro¬

podus and movable finger covered with small pits and

tubercles. Chelae of pereiopods II-III of equal length.

Pereiopods IV-V achelate. Lower margin of propodus of

pereiopod IV with a row of downeurved spines.

Abdominal appendages. Pleopods not preserved.

Discussion. Miinster (1839) described this species on

one specimen from Eichstàtt. Even though incomplete, thè

author identified some morphological characters, such as

thè rostrum with three strong supralateral teeth and very

elongate chelae of pereiopod I, in order to distinguish

this species from E. modestiforme (Schlotheim, 1822).

The study of thè originai specimen by Miinster confirms

thè validity of these characters as diagnostic, justifying

thè systematic validity of E. veltheimii (Miinster, 1839).

Moreover, thè presence of three characters not described

by Miinster, such as thè joining of postcervical and bran¬

chiocardiac grooves in their lower ends, thè upper margin

of propodus of pereiopod I with one strong distai spine

and thè telson with lateral margin spineless, are useful to

document thè systematic validity of this species and its

morphological differences from E. modestiforme.

Genus Palaeastacus Bell, 1850

Type species: Astacus sussexiensis Mantell, 1833

Palaeastacus fuciformis (Schlotheim, 1822)

Fig. 8, PI. I (fig. 3), PI. XI (fig. 5)

1822 - Macrourites fuciformis Schlotheim; p. 30, PI. 2 (fig. 2)

1827 - Astacus spinimanus Germar; p. 101, PI. 1 (fig. 3)

1829 -Astacus fuciformis Holl; p. 153

1839 - Glyphea fuciformis Miinster; p. 16, PI. 8 (figs. 1-2)

1839 - Glyphea intermedia Miinster; p. 17, PI. 8 (figs. 6-7)

1839 - Glyphea crassula Miinster; p. 17, PI. 8 (fig. 4) (not fig. 5)

1839 - Glyphea elongata Miinster; p. 18, PI. 8 (figs. 11-12) (not figs.

8-10)

1848 - Eryma fuciformis Bronn; p. 467

1852 -Astacus fuciformis Quenstedt; p. 268, PI. 20 (fig. 14)

1861 - Eryma elongata Mùnster; p. 358

1861 - Eryma fuciformis Schlotheim; p. 359

1862 - Eryma fuciformis (Schlotheim) in Oppel; p. 41, PI. 9 (figs. 2-6)

1862 -Eryma elongata Mùnster in Oppel; p. 37, PI. 8 (figs. 1-2)

not 1862 - Palaeastacus solitarius Oppel; p. 46, PI. 1 1 (fig. 4) = Pus-

tuli na suevica

1867 - Astacus fuciformis Quenstedt; p. 320, PI. 25 (fig. 14)

1885 - Astacus fuciformis Quenstedt; p. 409, PI. 32 (fig. 7)

1904 - Eryma fuciformis (Schlotheim) in Walther; p. 174

1925 - Eryma elongata Mùnster in Walther; p. 1 74

1925 - Enoploclytia fuciformis (Schlotheim) in Van Straelen; p. 285

not 1925 - Palaeastacus solitarius Oppel in Van Straelen; p. 289 = P

suevica

1928 - Eryma fuciformis (Schlotheim) in Beurlen; p. 164

1928 - Clytia elongata Mùnster in Beurlen; p. 170

1929 - Clytia sp. juv. in Glaessner; p. 1 19

10

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

1 929 - Eryma fuciformis (Schlotheim) in Glaessner; p. 1 54

not 1929 - Palaeastacus solitarius Oppel in Glaessner; p. 290 = P.

suevica

1961 - Eryma fuciformis (Schlotheim) in Kuhn; p. 22

not 1963 - Glyphea fuciformis Miinster in Miiller; p. 117, Text-fig. 157

(= Pustulina minuta)

1966 - Palaeastacus fuciformis (Schlotheim) in Forster; p. 130, Text-

fig. 25

1 968 - Palaeastacus fuciformis (Schlotheim) in Leich; p. 72

1 969 - Palaeastacus fuciformis (Schlotheim) in Malz; p. 299, Text-

fig. 8

not 1978 - Eryma fuciformis (Schlotheim) in Miiller; Text-fig. 148a (=

P minuta )

1994 - Palaeastacus fuciformis (Schlotheim) in Frickhinger; p. 122,

Text-figs. 218-219

2005b - Palaeastacus fuciformis (Schlotheim) in Schweigert & Ga-

rassino; p. 498

Diagnosis: carapace cylindrical laterally flattened;

dorsal margin dentate; cervical and branchiocardiac

grooves deep; gastro-orbital, postcervical, and ventral

grooves weak; antennal groove deep; hepatic notch and

muscle adductor testis weak; postcervical and branchio¬

cardiac grooves never joined; rostrum short with three

supralateral teeth; antennal spine well developed; pereio-

pods I-III chelate; pereiopod I larger and stronger than

pereiopods II-III; uropodal exopod with diaeresis.

Holotype: Macrourites fuciformis (= Palaeastacus

fuciformis ), MNHB K 36 MB. A. 251 (Schlotheim, 1822

- PI. 2, fig. 2; length of thè specimen: 3.5 cm), housed in

thè Museum fur Naturkunde der Humboldt-Universitàt in

Berlin.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - lower Tithonian).

Type locality: Solnhofen (?).

Material: Oppel (1862) reported 70 specimens

belonging to this species, so divided: 22 from thè Reden-

bacher collection, three from thè collection of Prof. O.

Fraas in Stuttgart (SMNS, but not traceable) and 45 from

thè collection of thè Bayerische Staatssammlung fur Pala-

ontologie und Geologie in Munich. We studied of this

originai sample thè holotype by Schlotheim, housed in

thè Museum fùr Naturkunde der Humboldt-Universitàt

in Berlin, and nine specimens, housed in thè Bayerische

Staatssammlung fur Palàontologie und Geologie in

Munich in which we identifìed five specimens, illustrated

by Miinster (1839) and four specimens, illustrated by

Oppel (1862). Two specimens (BSPG AS I 620, AS VI

14 a-b) were identifìed as belonging to originai collec-

tions thanks to thè 20th century labels. However, it was

impossible to establish if they were studied by one of thè

above-mentioned authors because thè specimens were not

illustrated. We report thè list of thè specimens, studied by

Miinster and Oppel:

BSPG AS VII 183 - Miinster (1839), PI. 8 (fig. 2)

- length of thè specimen: 4.5 cm

BSPG AS VII 184 - Miinster (1839), PI. 8 (fig. 4)

- length of thè specimen: 3.5 cm

BSPG AS VII 185 - Miinster (1839), PI. 8 (fig. 7)

- length of thè specimen: 4.5 cm

BSPG AS VII 192 - Miinster (1839), PI. 8 (fig. 11)

- length of thè specimen: 1.5 cm

BSPG AS VI 11 - Oppel (1862), PI. 9 (figs. 5-6)

- length of thè specimen: 4 cm

BSPG AS VI 12 - Oppel (1862), PI. 9 (fig. 4)

- length of thè specimen: 3.5 cm

BSPG AS VI 13 - Oppel (1862), PI. 9 (figs. 2-3)

- length of thè specimen: 3.5 cm

BSPG AS VI 18 - Oppel (1862), PI. 8 (fig. 1)

- length of thè specimen: 1 .5 cm

BSPG AS Vili 83 - Oppel (1862), PI. 8 (fig. 2)

- incomplete specimen

Description. Small-sized erymid with strongly spiny

and tuberculate exoskeleton.

Carapace. Carapace cylindrical laterally flattened.

Dorsal margin dentate. Rostrum short with three supralat¬

eral teeth. Cervical and branchiocardiac grooves deep.

Gastro-orbital, postcervical, and ventral grooves weak.

Antennal groove deep. Hepatic notch and muscle adduc¬

tor testis weak. Antennal spine well developed. Ocular

incision large and deep. Spines short and strong located

in gastric, antennal, and in thè upper part of branchial

regions. Spines arranged in two rows in thè gastric region,

increasing in size toward thè ocular incision. Tubercles in

thè pterygostomial and in thè median and lower parts of

branchial regions.

Abdomen. Somites I-VI subrectangular of equal

length and with smooth tergal surface. Telson subtrian-

gular with pointed distai extremity, deep median longi-

tudinal groove, marked by two parallel rows of small

spines, two oblique weak carinae and three strong spines,

increasing in size from thè proximal to thè distai one, in

thè median part of lateral margins. Protopod subrectangu¬

lar. Uropodal endopod with a strong median longitudinal

carina. Uropodal exopod with a strong median longitu¬

dinal carina, two distai spines on thè outer margin (one

longer and one shorter), rounded diaeresis with serrate

upper margin having one longer median spine and tuber¬

culate lower margin.

Cephalic appendages. Eyestalk short. 3rd maxilliped

elongate with first proximal article having two strong

distai spines and second article with one strong distai

spine, all located on thè lower margin. Antennular arti-

cles thin of which thè distai one carries two elongate

multiarticulate flagella of equal length. Antennal articles

strong and stout of which thè distai one carries a strong

elongate multiarticulate flagellum, twice thè body length.

Scaphocerite elongate and triangular with serrate lower

margin and with basicerite having a strong distai spine.

Thoracic appendages. Pereiopod I with short and stout

chela. Movable and fixed fingers of equal length with

curved distai extremity. Fixed fìnger with strongly dentate

inner margin. Inner margin of movable finger edentate.

Upper margin of merus with a row of strong spines,

increasing in length toward thè articolation with carpus.

Lower margin of merus with a row of strong spines of

equal length. Upper margin of carpus, propodus and mov¬

able finger with a row of strong spines of equal length.

Lower margin of carpus, propodus and fixed fìnger with

a row of small spines of equal length. Surface of merus,

carpus, propodus and movable finger covered with tuber¬

cles and strong spines arranged in parallel rows. Chela of

pereiopod III longer than chela of pereiopod II. Chelae

of pereiopods II-III with a strong spine in thè articulation

between movable and fixed fingers. Pereiopods IV longer

and stronger than pereiopod V, both achelate. Lower

margin of propodus of pereiopod IV with a row of down-

curved spines.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

11

Abdominal appendages. Pleopods with an unseg-

mented peduncle which carries two elongate multiarticu-

late flagella.

Discussion. Forster (1966) reviewed this genus,

describing seven species. Today, Palaeastacus Bell,

1850, known from thè Lower Jurassic (Sinemurian) to

thè Paleocene of Europe (France, Switzerland, Germany

and England), Antarctic Peninsula, United States, and

Australia, includes 18 species. Eight from thè Jurassic:

P. falsarli (Dumortier, 1867) (France - Sinemurian), P.

argoviensis Forster & Rieber, 1982 (Switzerland - Aale-

nian), P. spinosus (Etallon, 1861) (France - Aalenian),

P edwardsi Etallon, 1861 (France - Callovian), P. fuci-

formis (Schlotheim, 1822) (Germany - Tithonian), P.

rothgaengerae Schweigert & Ròper, 2001 (Germany

- Kimmeridgian), P. poeschli Schweigert & Ròper, 2001

(Germany - Tithonian), and 7? decoratus (Frentzen, 1937)

(Germany - Pliensbachian). Nine from thè Cretaceous: P.

foersteri Taylor, 1979 (Antarctic Peninsula - Aptian), P.

scaber (Bell, 1 863) (England - Albian), P. terraereginae

(Etheridge, 1914) (Australia - Albian), P. sussexiensis

(Mantell, 1833) (England - Cenomanian), P. trìsulcatus

Schweitzer & Feldmann, 2001 (United States - Ceno¬

manian), P triglyptus (Stenzel, 1945) (United States

- Coniacian), P kimzeyì Rathbun, 1935 (United States

- Campanian), P. dixoni Bell (1850) (England - Chalk

Formation), and P. macrodactylus Bell, 1850 (England

- Chalk Formation). One from thè Tertiary: P selmaensis

Rathbun, 1935 (United States - Paleocene).

Genus Pustulina Quenstedt, 1857

Type species: Pustulina suevica Quenstedt, 1857

Pustulina minuta (Schlotheim, 1822)

Fig. 8, PI. II (fig. 1), PI. XI (fìg. 6), PI. XII (fìgs. 1, 2)

1822 - Macrourites minutus Schlotheim; p. 28, PI. 3 (fig. 3)

1839 - Glyphea minuta Mtinster; p. 20, PI. 9 (fìgs. 8-10)

1839 - Glyphea verrucosa Miinster; p. 21, PI. 9 (fig. 11) (not fig. 12 =

lectotype of Eryma verrucosa)

1 861 - Eryma minuta (Schlotheim) in Oppel; p. 359

1862 - Eryma minuta (Schlotheim) in Oppel; p. 39, PI. 8 (figs. 6-8)

1904 - Eryma minuta (Schlotheim) in Walther; p. 174

1925 - Enoploclytia minuta (Schlotheim) in Van Straelen; p. 284

1 925 - Enoploclytia fuciformis (Schlotheim) in Van Straelen; p. 285 (pars)

1928 - Eryma minuta (Schlotheim) in Beurlen; p. 164

1929 - Eryma fuciformis (Schlotheim) in Glaessner; p. 154 (pars)

1929 -Eryma minuta (Schlotheim) in Glaessner; p. 156

1961 - Eryma minuta (Schlotheim) in Kuhn; p. 22

1966 - Phlyctisoma minuta (Schlotheim) in Forster; p. 142, PI. 1 8 (fig. 9)

1969 - Phlyctisoma minuta (Schlotheim) in Malz; p. 300, Text-fig. 9

not 1994 - Phlyctisoma minuta (Schlotheim) in Frickhinger; p. 126,

Fig. 224 = Palaeastacus sp.

Diagnosis: carapace cylindrical laterally flattened; ros-

trum short with three small supralateral teeth; gastro-orbital

and cervical grooves deep; postcervical groove strongly

developed reaching hepatic groove; branchiocardiac

groove extremely small; antennal spine well developed;

pereiopods I-III chelate; pereiopod I larger and stronger

than pereiopods II-III; uropodal exopod with diaeresis.

Holotype: Macrourites minutus (= Pustulina minuta),

MNHB K 41 MB (Schlotheim, 1822 - PI. 3, fig. 3;

length of thè specimen: 2 cm), housed in thè Museum fur

Naturkunde der Humboldt-Universitàt in Berlin.

Our studies revealed that one thè syntypes of Glyphea

verrucosa from Eichstàtt and figured by Mtinster (1839

- PI. 9, fig. 11) belongs to Pustulina minuta, as already

recognized by Oppel (1862)

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Eichstàtt (?).

Material: Oppel (1862) reported 44 specimens

belonging to this species, so divided: 13 from thè Reden-

bacher collection, one from collection of thè Museum fur

Naturkunde der Humboldt-Universitàt in Berlin and 30

from thè collection of thè Bayerische Staatssammlung

fur Palàontologie und Geologie in Munich. We studied of

this originai sample thè holotype by Schlotheim, housed

in thè Museum far Naturkunde der Humboldt-Universitàt

in Berlin, and four specimens, housed in Bayerische

Staatssammlung fur Palàontologie und Geologie in

Munich in which we identifìed two specimens, illus-

trated by Miinster (1839) and two specimens, illustrated

by Oppel (1862). One specimen (BSPG AS I 619) was

identifìed as belonging to originai collections thanks to

thè 19th century labels. However, it was impossible to

establish if it was studied by one of thè above-mentioned

authors because it was not illustrated. We report thè list of

thè specimens, studied by Miinster and Oppel:

BSPG AS VII 180 - Miinster (1839), PI. 9 (fìg. 10)

- length of thè specimen: 3 cm

BSPG AS VII 181 - Miinster (1839), PI. 9 (fig. 11)

- length of thè specimen: 3 cm

BSPG AS VI 10 - Oppel (1862), PI. 8 (fig. 8)

- length of thè specimen: 3 cm

BSPG AS Vili 78 - Oppel (1862), PI. 8 (fig. 7)

- length of thè specimen: 4 cm

Oppel (1862) reported one specimen, an isolated

pereiopod I, which he kept in Eryma verrucosa (Miinster,

1839), hence taking that specimen as thè lectotype of this

taxon. It was housed in thè Bayerische Staatssammlung

fur Palàontologie und Geologie in Munich where, today,

it is stili preserved.

BSPG AS VII 182 -Miinster (1839), PI. 9 (fìg. 12)

BSPG AS VII 1 82 - Oppel ( 1 862), PI. 8 (fig. 5)

- incomplete specimen

One specimen (n. cat. 0314 - Wulf collection) was

used to describe thè tail fan of this species, poorly pre¬

served in thè originai sample.

Description. Small-sized erymid with densely tuber-

culate exoskeleton.

Carapace. Carapace cylindrical laterally flattened. Ros-

trum short with three small supralateral teeth. Cervical and

gastro-orbital grooves deep. Postcervical groove strongly

developed reaching hepatic groove. Branchiocardiac

groove extremely small. Antennal, hepatic, and ventral

grooves weak. Antennal spine well developed. Ocular inci-

sion small. Dorsal surface of carapace densely tuberculate.

Abdomen. Somites I-V subrectangular of equal length

and with finely tuberculate tergal surface. Somite VI

subsquare. Telson subtriangular with weak median lon-

gitudinal groove, two pairs of small tubercles in proximal

part of dorsal surface, one pair of small distai spine on

lateral margin, and rounded distai extremity. Protopod

subrectangular. Uropodal endopod with strong longitudi-

12

ALESSANDRO GARASSfNO & GUNTER SCHWEIGERT

nal median carina. Uropodal exopod with a weak longi-

tudinal median carina, two different distai spines on thè

outer margin (one longer and one shorter) and rounded

diaeresis with serrate upper margin.

Cephalic appendages. Eyestalk short. 3rd maxilliped

elongate with thè first article having a strong distai spine

in thè lower margin. Antennular articles thin of which thè

distai one carries two elongate multiarticulate flagella of

equal length. Antennal articles strong and stout of which

thè distai one carries a strong elongate multiarticulate

flagellum, twice thè body length.

Thoracic appendages. Pereiopod I with very short

and stout chela. Movable finger longer than fixed fìnger

with curved distai extremity. Inner margins of movable

and fixed fingers edentate. Surface of merus, carpus,

propodus, movable, and fixed fingers covered with strong

tubercles. Chelae of pereiopods II-III of equal length.

Pereiopods IV-V achelate. Lower margin of propodus of

pereiopod IV with a row of downcurved spines.

Abdominal appendages. Pleopods not preserved.

Discussion. Phlyctisoma Bell, 1863, now synonym

of Pustulina Quenstedt, 1857, after thè review by Sch-

weigert et al. (2000), was studied by Forster (1966) who

described seven species. Today, Pustulina , known from

thè Lower Jurassic (Sinemurian) to thè Upper Cretaceous

(Campanian - Maastrichtian) of Europe (Germany, Eng-

land, France and Italy), Madagascar and Canada, includes

12 species. Seven from thè Jurassic: P. sinemuriana (Gar-

assino, 1996) (Italy - Sinemurian), P. elegans (Méchin,

1901) (France - Bajocian), P. calloviensis (Forster,

1966) (Germany - Callovian), P. perroni (Etallon, 1861)

(France - Oxfordian), P pseudobabeaui (Dollfus, 1863)

(France - Kimmeridgian), P. suevica Quenstedt, 1857

(Germany - Kimmeridgian), and P. minuta (Schlotheim,

1822) (Germany - Tithonian). Five from thè Cretaceous:

P. spinulata (Secretan, 1964) (Madagascar - Valangin-

ian-Hauterivian), P. tuberculata (Bell, 1863) (England

- Albian), P. granulata (Bell, 1863) (England - Albian),

P. scabra (Bell, 1863) (England - Albian), and P. dawsoni

(Woodward, 1900) (Canada - Upper Cretaceous).

Miinster (1839) described Glyphea verrucosa on one

poorly preserved specimen (fìgured in PI. 9, fig. 11) and

one isolated complete chela. Only thè latter was kept

in this species by Oppel (1862), so that this chela has

become thè lectotype of Glyphea verrucosa Miinster,

1839. Forster (1966) discussed thè systematic position of

E. verrucosa (Miinster, 1839), rejecting thè ascription of

this species to Pustulina minuta (Schlotheim, 1822), for

thè different omamentation of thè chela. Moreover, thè

author, comparing E. verrucosa with E. boloniensis Sau-

vage, 1891 (France - Kimmeridgian) and E. mosquensìs

Lahusen, 1 894 (Russia - Portlandian), both preserved as

incomplete chelae, pointed out thè existence of a mor-

phological affinity between thè propodus of thè German

species and thè other two. However, thè latter view was

based on a large chela in thè collection of thè Naturkun-

demuseum in Dresden which Forster (1966) assigned to

thè same species. The latter, unfigured specimen of which

a plaster cast exists in thè Munich collection, clearly

belongs to Erymastacus sp. in respect of its very large

fingers showing occludent margins with well-developed,

regularily spaced teeth. The lectotype of Glyphea ver¬

rucosa Miinster, 1839, differs from Pustulina minuta by

thè presence of a densely pustulate omamentation of thè

propodus and its more straight fingers. To date no other

specimen of this rare taxon could be identified in collec-

tions, so that thè generic assignment is uncertain.

Family Protastacidae Albrecht, 1983

Genus Pseudastacus Oppel, 1861

Type species: Bolina pustulosa Miinster, 1839

Pseudastacus pustulosus (Miinster, 1839) (c^)

Fig. 8, PI. II (fig. 2), PI. XII (figs. 3, 4)

1839 - Bolina pustulosa Miinster; p. 23, PI. 9 (fig. 13)

1840 - Alvis octopus Miinster; p. 20, PI. 1 (fig. 1)

1861 - Pseudastacus pustulosus (Miinster) in Oppel; p. 360

1861 - Pseudastacus Miinsteri Oppel; p. 360, nov. syn. ($)

1 862 - Pseudastacus pustulosus (Miinster) in Oppel; p. 44, PI. 10 (figs.

4-5), PI. 11 (fig. 1)

1 862 - Pseudastacus Miinsteri Oppel; p. 45, PI. 1 1 (fig. 2)

1904 - Pseudastacus pustulosus (Miinster) in Walther; p. 174

1904 - Pseudastacus Miinsteri Oppel in Walther; p. 174

1925 - Pseudastacus pustulosus (Miinster) in Van Straelen; p. 292,

Text-fig. 133

1 929 - Pseudastacus pustulosus (Miinster) in Glaessner; p. 35 1

1 969 - Pseudastacus pustulosus (Miinster) in Glaessner; R 460, Text-

fig. 267/1

1994 - Pseudastacus pustulosus (Miinster) in Frickhinger; p. 26, Fig.

225

Diagnosis: carapace cylindrical laterally flattened;

cervical, gastro-orbital, antennal, and hepatic grooves

deep; postcervical groove weak; branchiocardiac groove

absent; dorsal suture absent; rostrum long with three

strong supralateral teeth; antennal spine weak; pereiopods

I-III chelate; pereiopod I larger and stronger than pereio¬

pods II-III; uropodal exopod with diaeresis.

Lectotype: BSPG AS I 670 (Oppel, 1 862 - PI. 1 0, fig.

4), housed in thè Bayerische Staatssammlung fur Palàon-

tologie und Geologie in Munich.

Our studies revealed that Pseudastacus muensteri from

Solnhofen and fìgured by Oppel ( 1 862 - PI. 11, fig. 2 a-b)

is synonym with Pseudastacus pustulosus , as discussed

below. This specimen, BSPG AS I 672, is thè holotype

of Oppel ’s species, housed in thè Bayerische Staatssa¬

mmlung ftir Palaontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported ten specimens

belonging to this species, so divided: one from thè Reden-

bacher collection and nine from thè collection of thè Bay¬

erische Staatssammlung fur Palaontologie und Geologie

in Munich. We studied of this originai sample four speci¬

mens, housed in thè Bayerische Staatssammlung fur Pala-

ontologie und Geologie in Munich in which we identified

one specimen, illustrated by Miinster (1839) and three

specimens, illustrated by Oppel (1862). Seven specimens

were identified as belonging to originai collections thanks

to thè 19^ century labels; two to Miinster collection, four

to Oppel collection and one unidentified. Finally, thè

specimen BSPG AS VII 323, belonging to Miinster ’s col¬

lection was not illustrated by thè author. We report thè list

of thè specimens, studied by Miinster and Oppel:

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART 1

13

BSPG AS VII 321 - Miinster (1839), PI. 9 (fig. 13)

- length of thè specimen: 6 cm

BSPG AS I 669 - Oppel (1862), PI. 10 (fig. 5)

- length of thè specimen: 6 cm

BSPG AS I 670 - Oppel (1862), PI. 10 (fig. 4)

- length of thè specimen: 6 cm

BSPG AS I 671 - Oppel (1862), PI. 11 (fig. 1)

- length of thè specimen: 6 cm

Oppel (1862) reported one specimen which he

assigned to Pseudastacus muensteri Oppel, 1862, housed

in thè Bayerische Staatssammlung fur Palàontologie und

Geologie in Munich where, today, it is stili preserved.

BSPG AS I 672 - Oppel (1862), PI. 11 (fig. 2 a-b)

- length of thè specimen: 4 cm

One well-preserved specimen (n. cat. SMNS 65480

- Koschny collection) was used to describe some mor¬

phological characters of thè carapace, poorly preserved in

thè originai specimens.

Description. Median-sized protoastacid with strongly

tuberculate exoskeleton.

Carapace. Carapace cylindrical laterally flattened.

Dorsal margin straight. Dorsal suture absent. Rostrum

long with three strong supralateral teeth. Cervical, gastro-

orbital, antennal, and hepatic grooves deep. Postcervical

groove weak. Branchiocardiac groove absent. Antennal

spine weak. Ocular incision large and deep.

Abdomen. Somites I-VI subrectangular of equal

length with slightly tuberculate tergal surface. Telson

subrectangular with rounded distai extremity and one

strong distai spine on thè lateral margin. A kind of trans¬

verse fold breaks telson into two almost identical parts:

rigid and tuberculate upper part and flexible lower part.

Protopod subrectangular. Uropodal endopod with a

strong longitudinal median carina. Uropodal exopod with

a strong longitudinal median carina, strong distai spine

on thè outer margin, and rounded diaeresis with serrate

upper margin.

Cephalic appendages. Eyestalk short. 3rd maxilliped

elongate and spineless. Antennular articles thin of which

thè distai one carries two elongate multiarticulate flagella

of equal length. Antennal articles strong and elongate of

which thè distai one carries a strong elongate multiarticu¬

late flagellum, twice thè body length. Scaphocerite strong,

elongate, and triangular with pointed distai extremity.

Thoracic appendages. Pereiopod I with short and stout

chela. Movable and fixed fingers of equal length with

curved distai extremity. Serrate inner margins of movable

and fixed fingers. Lower margin of propodus with a row

of small spines. Dorsal surface of merus with a longitu¬

dinal median row of strong tubercles. Lower margins of

carpus and merus with small spines. Strong median spine

on articulate margins of merus and carpus. Surface of

merus, carpus, propodus, and movable finger covered

with small tubercles. Chela of pereiopod II smaller than

chela of pereiopod III. Pereiopods IV-V achelate.

Abdominal appendages. Pleopods with an unseg-

mented peduncle which carries two elongate multiarticu¬

late flagella.

Discussion. Pseudastacus Oppel, 1861, was included

for long time in thè family Nephropidae Dana, 1852,

as reported by Glaessner (1969) and Chong & Forster

(1976). Albrecht (1983) reviewed this genus, includ-

ing it in thè new family Protastacidae Albrecht, 1983

(infraorder Astacidea Latreille, 1802). The author justi-

fied thè institution of this family because it represents a

transitional grade between thè erymids and crayfishes.

Tshudy & Babcock (1997) included Pseudastacus in their

newly erected family Chilenophoberidae, but their analy-

sis is not founded by a morphological review of thè type

material but only by line drawings in literature.

Today, thè family Protastacidae includes two genera:

Pseudastacus Oppel, 1861, with two species, P. pustulo-

sus Miinster, 1839, and P. minor Fraas, 1878 (Germany

- Tithonian), and Protoastacus Albrecht, 1983, with P.

politus (Schliiter, 1 868) (Germany - Upper Cretaceous).

The holotype of P. dubertreti Roger, 1946 (n. cat. B.

18892) (Lebanon - Cenomanian) housed in thè Muséum

national d’Histoire naturelle in Paris, was recently exam-

ined by one of thè authors (A. Garassino). Its study has

pointed out that thè elongate rostrum with a row of dorsal

teeth and thè annulated carpus of pereiopods II-III are two

characters typical of Carpopenaeus callirostris Glaessner,

1945. So P dubertreti must be considered as synonym

with C. callirostris.

Oppel (1862), describing P muensteri, pointed out

that thè small size of thè body and thè longer and sten¬

der pereiopod I distinguished this new species from P.

pustulosus (Miinster, 1839). Apart from this very sig-

nificant character, thè specimen belonging to P. muensteri

exhibits thè same morphological characters of thè type

species, such as deep cervical, gastro-orbital, antennal

and hepatic grooves, weak postcervical groove, branchio¬

cardiac groove absent, dorsal suture absent, long rostrum

with three strong supralateral teeth, weak antennal spine

and uropodal exopod with diaeresis. After these remark-

able characters in common between P pustulosus and

P muensteri, we suppose these forms represent sexual

dimorphs of a single species. In fact, in Recent glypheid

Neoglyphea inopinata, and other fossil glypheids, another

group of decapods, thè sexual dimorphism also affects

thè lengths of pereiopod I and its articles, with thè longer

ones are developed in thè female (Forest & de Saint Lau¬

rent, 1981, 1989). Thus we suppose P. pustulosus is thè

male and P muensteri thè female form (see also thè sexual

dimorphs in Cycleryon Glaessner, 1965, and Mecochirus

Germar, 1827, in this paper).

Family Stenochiridae Beurlen, 1930

Discussion. Beurlen (1930) established thè family

Stenochiridae that includes Stenochirus Oppel, 1861,

without defining its diagnostic characters. The author in

fact based this species mainly on thè characters of thè

monotypic Stenochirus suevicus Quenstedt, 1867 (Ger¬

many - Aalenian), despite S. angustus Miinster, 1839, thè

type species of thè genus. Stenochirus suevicus became

later type species of thè genus Palaeophoberus Glaessner,

1932. By thè study of two very well preserved specimens

(n. cat. 9706, 9836 - Wulf collection), it was possible to

frame for thè first time thè diagnosis of this family. Thus

we do not follow thè ascription of Stenochirus in Chileno¬

phoberidae by Tshudy & Babcock (1997).

Diagnosis: carapace cylindrical laterally flattened;

rostrum long with four supralateral teeth; two dentate car-

inae in antennal region; cervical groove deep; branchio¬

cardiac groove weak ; pereiopods l-III chelate; pereiopod

14

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

I larger and stronger than pereiopods II-III; pereiopods II-

V weak with slender and elongate segments; pereiopods

II-III weak hearing very small chelae; somite I remark-

ably smaller than thè others; expanded pleura of somite

II not overlapping that of somite I; uropodal exopod with

diaeresis.

Genus Stenochirus Oppel, 1861

Type species: Bolina angusta Miinster, 1839

Solnhofen species: Stenochirus angustus (Miinster,

1839); Stenochirus mayeri Oppel, 1862

Stenochirus angustus (Miinster, 1839)

Fig. 8, PI. Ili (fig. 1), PI. XII (figs. 5, 6)

1839 - Bolina angusta Miinster; p. 24, PI. 9 (fig. 14)

1853 - Bolina angusta Miinster in Frischmann; p. 26

1862 - Stenochirus angustus (Miinster) in Oppel; p. 20, PI. 4 (figs.

3-4)

1904 - Stenochirus angustus (Miinster) in Walther; p. 174

1925 - Stenochirus angustus (Miinster) in Van Straelen; p. 131

1929 - Stenochirus angustus (Miinster) in Glaessner; p. 380

1994 - Stenochirus angustus (Miinster) in Frickhinger; p. 127, Fig.

226

1999 - Stenochirus angustus (Munster) in Frickhinger; p. 44, Fig. 71

2005b - Stenochirus angustus (Miinster) in Schweigert & Garassino;

p. 498

The poor state of preservation of thè originai specimen

by Oppel has led to thè examination of new specimens

from private collections in order to expand thè morpho-

logical description of Stenochirus angustus. Two speci¬

mens (n. cat. 9706, 9836 - Wulf collection), preserved

in lateral and dorsal views allowed recognition of many

important morphological characters in order to give for

thè first time a detailed description of this species and thè

systematic position of thè genus into thè family Astacidae

Latreille, 1802.

Diagnosis: as for thè genus.

Holotype by monotypy: BSPG AS VII 322 (Munster,

1839 - PI. 9, fig. 14), housed in thè Bayerische Staatssa-

mmlung fur Palàontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Flonheim near Solnhofen.

Material: Oppel (1862) reported one specimen, thè

holotype of Munster, ascribed to this species, and housed

in thè Bayerische Staatssammlung tur Palàontologie und

Geologie in Munich where today it is stili preserved.

BSPG AS VII 322 - Munster (1839), PI. 9 (fig. 14)

BSPG AS VII 322 - Oppel (1862), PI. 4 (figs. 3-4)

- length of thè specimen: 3 cm

Description. Median-sized astacid with smooth

exoskeleton.

Carapace. Carapace cylindrical laterally flattened.

Rostrum long with four small supralateral teeth directed

forward. Cervical groove median and deep. Branchiocar-

diac groove weak. Two dentate carinae in antennal region.

Posterior margin slightly convex. Ocular incision small,

and narrow. Dorsal surface of carapace smooth.

Abdomen. Somite I remarkably smaller than thè

others. Somites II-VI subrectangular of equal length,

rounded pleurae, and with smooth tergal surface. Pleura

of somite II subrounded overlapping that of somite III.

Telson subrectangular with a weak median longitudinal

groove, two pairs of small median longitudinal spines on

dorsal surface, and two oblique carinae originating in thè

proximal part of dorsal surface terminating with a small

distai spine on thè lateral margin. Protopod subrectangu¬

lar. Uropodal endopod with a weak longitudinal median

carina. Uropodal exopod with a weak longitudinal median

carina, one small distai spine on thè outer margin, and

rounded diaeresis.

Cephalic appendages. Eyestalk short. Antennular

articles thin of which thè distai one carries two elon¬

gate multiarticulate flagella of equal length. Antennal

articles strong and stout of which thè distai one carries

a strong elongate multiarticulate flagellum of thè same

body length. Scaphocerite triangular with pointed distai

extremity.

Thoracic appendages. Pereiopod I with very elongate

and slender chela. Movable and fixed fìngers of thè same

length with curved distai extremity. Inner margins of

movable and fixed fingers with row of thin and elongate

teeth of thè same length directed forward. Upper margin

of merus with a row of small tubercles. Dorsal surface of

propodus with a strong median longitudinal carina. Upper

and lower margins of merus and carpus of pereiopod I

with a row of strong spines directed forward. Pereiopods

II-V weak hearing elongate and slender articles. Pereio¬

pods II-III with remarkably very small chelae. Pereiopods

IV- V achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. The systematic position of Stenochirus

Oppel, 1861, was uncertain for a long time. Beurlen

(1928, 1930) established thè subfamily Stenochirinae

originally and thè family Stenochiridae later, justifying

it on thè basis of morphological characters that were

different from thè other astacideans. Glaessner (1929,

1969) included Stenochirus in thè family Erymidae Van

Straelen, 1924, originally and later he did not include this

genus in any known family of thè infraorder Astacidea

Latreille, 1803.

The study of thè originai and new specimens permit-

ted recognition of some morphological characters, such

as weak pereiopods II-V hearing slender and elongate

articles, pereiopods II-III with remarkably small chelae,

somite I remarkably smaller than thè others, and sub¬

rounded pleura of somite II overlapping that of somite

III, that are typical of Stenochirus and are not observ-

able in thè other genera within thè infraorder Astacidea

in order to attest thè originai institution of thè family by

Beurlen.

Moreover, among thè genera belonging to thè

infraorder Astacidea, thè above-mentioned characters are

also observable in Uncina Quenstedt, 1851 (family Unci-

nidae Beurlen, 1928) recently reviewed by Schweigert et

al. (2003), even though this genus differs from Stenochi¬

rus on thè basis of thè different shape and morphology of

thè chela of pereiopod I.

Today, thè family Stenochiridae Beurlen, 1930, previ-

ously known only in thè Upper Jurassic (Tithonian) of

Germany, includes three species, S. angustus Munster,

1839, S. mayeri Oppel, 1862, and S. vahldieki Sch¬

weigert, Garassino & Riou, 2006, from thè Callovian of

France (Schweigert et al., 2006).

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART

15

Stenochirus mayeri Oppel, 1 862

Fig. 2, PI. XIII (figs. 1,2)

1862 - Stenochirus Mayeri Oppel; p. 20, PI. 4 (figs. 1-2)

1904 - Stenochirus Meyeri Oppel in Walther; p. 175

1925 - Stenochirus Meyeri Oppel in Van Straelen; p. 131

1929 - Stenochirus Meyeri Oppel in Glaessner; p. 380

1969 - Stenochirus meyeri Oppel in Glaessner; R 460, Text-fig. 267/2

1994 - Stenochirus meyeri Oppel in Frickhinger; p. 127, Fig. 227

Lectotype: BSPG AS I 673 (Oppel, 1862 - PI. 4, figs.

1-2), housed in thè Bayerische Staatssammlung tur Pala-

ontologie und Geologie in Munich.

Stratigraphic range; Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Remark: The correct originai spelling of this taxon is

“mayeri". An according to thè originai hand-written label

by Oppel this species was dedicated to Charles Mayer-

Eymar, not to Hermann v. Meyer.

Material: Oppel (1862) reported one specimen,

ascribed to this species, and housed in thè Bayerische

Staatssammlung fur Palàontologie und Geologie in

Munich where, today, it is stili preserved.

BSPG AS I 673 - Oppel (1862), PI. 4 (figs. 1-2)

- length of thè specimen: 3 cm

Discussion. This species was described by Oppel

(1862) on one specimen in ventral view. Even though,

thè poor state of preservation of thè originai specimen

made difficult to observe thè morphological characters

of Stenochirus Oppel, 1861, pereiopod II with remark-

ably small chela, pleura of somite II overlapping that

Fig. 2 - Stenochirus mayeri Oppel, 1862, chela of pereiopod I, line

drawing (F. Fogliazza).

of somite III, and uropodal exopod with small spine in

thè distai part of lateral margin are enough to validate

its ascription to this genus. However, it is difficult to

develop a morphological description of this species

because it is preserved in ventral view. We agree with

Oppel in regard to its systematic validity because thè

shape of thè chela of pereiopod I is different from that of

S. angustus Miinster, 1839 (Fig. 2). In fact, Oppel’s spe¬

cies exhibits a slender, slightly curved propodus of thè

chela of pereiopod I with thè movable finger longer than

fixed fìnger. Miinster’s species has a stout and straight

propodus of chela of pereiopod I with movable and fixed

fìngers of thè same length. In both species, thè inner

margins of movable and fixed fìngers have a row of thin

and elongate teeth of thè same length directed forward.

Only thè discovery of well preserved specimens will

make possible a complete morphological description of

this species.

Family Glypheidae Winckler, 1883

Genus Glyphea v. Meyer, 1835

Type species: Palinurus regleyanus Desmarest, 1822

Solnhofen species: Glyphea pseudoscyllarus (Schlot-

heim, 1822); Glyphea tenuis Oppel, 1860

Glyphea pseudoscyllarus (Schlotheim, 1822)

Fig. 9, PI. Ili (fig. 2), PI. XIII (fig. 3)

1822 - Macrourites pseudoscyllarus Schlotheim; p. 36, PI. 12 (fig.

5)

1829 - Scyllarus dubius Holl; p. 151

1839 - Orphnea pseduoscyllarus Miinster; p. 39, PI. 14 (figs. 1-2)

1839 - Orphnea striata Miinster; p. 40, PI. 14 (fig. 3)

1839 - Orphnea laevigata Miinster; p. 41, PI. 14 (fig. 5)

1839 - Orphnea pygmaea Miinster; p. 42, PI. 14 (fig. 6)

1839 - Brisa dubia Miinster; p. 46, PI. 15 (figs. 4-5?)

1839 - Brisa lucida Miinster; p. 46, PI. 15 (fig. 3)

1853 - Orphnea pseudoscyllarus Miinster in Frischmann; p. 29

1853 - Orphnea striata Miinster in Frischmann; p. 29

1860 - Glyphea pseudoscyllarus (Schlotheim) in Oppel; p. 1 10

1862 - Glyphea pseudoscyllarus (Schlotheim) in Oppel; p. 72, PI. 18

(fig. 2), PI. 19 (figs. 1-5)

1883 - Glyphea pseudoscyllarus (Schlotheim) in Winckler; p. 26, Figs.

4,6

1904 - Glyphea pseudoscyllarus (Schlotheim) in Walther; p. 174

1922 - Glyphea pseudoscyllarus (Schlotheim) in Van Straelen; p.

1224-1226

1925 - Glyphea pseudoscyllarus (Schlotheim) in Van Straelen; p. 192,

Text-fig. 94

1928 — Glyphea pseudoscyllarus (Schlotheim) in Beurlen; p. 140

1929 - Glyphea pseudoscyllarus (Schlotheim) in Glaessner; p. 189

1969 - Glyphea pseudoscyllarus (Schlotheim) in Glaessner; R 463

not 1994 - Glyphea pseudoscyllarus (Schlotheim) in Frickhinger; p.

120, Fig. 207 = G. tenuis

2001 - Glyphea pseudoscyllarus (Schlotheim) in Dietl & Schweigert;

p. 55, Text-fig. 90

Diagnosis: carapace cylindrical laterally flattened;

rostrum very short with curved distai extremity; cervical

and branchiocardiac grooves deep; postcervical groove

weak, joined to branchiocardiac groove near their lower

16

ALESSANDRO GARASSINO & GÙNTER SCHWEIGERT

ends; antennal, hepatic, and ventral grooves weak; three/

four carinae tuberculate extend parallel in gastric and

antennal regions; pereiopod I subchelate; pereiopods II-V

achelate; uropodal exopod with diaeresis.

Holotype: Macrourites pseudoscyllarus (= Glyphea

pseudoscyllarus), MNHB K 43 MB. A. 255 (Schlotheim,

1822 - PI. 12, fig. 5; length of thè specimen: 5 cm),

housed in thè Museum fur Naturkunde der Humboldt-

Universitàt in Berlin.

Stratigraphic range: Upper Jurassic (upper Kimmer-

digian - lower Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported 50 specimens

belonging to this species, so divided: 16 from thè

Redenbacher collection and 34 from thè collection

of thè Bayerische Staatssammlung fur Palàontologie

und Geologie in Munich. We studied thè holotype by

Schlotheim, housed in thè Museum fur Naturkunde der

Humboldt-Universitat in Berlin, and nine specimens of

this originai sample, housed in thè Bayerische Staatssa¬

mmlung fur Palaontologie und Geologie in Munich

in which we identified one specimen, illustrated by

Munster (1839) and Oppel (1862) and seven specimens,

illustrated by Oppel (1862). Six specimens without

catalogue numbers were identified as belonging to

originai collections based upon thè 19th century labels.

We report thè list of thè specimens, studied by Munster

and Oppel:

BSPG AS VII 730 - Munster (1839), PI. 14 (fig. 2)

BSPG AS VII 730 - Oppel (1862), PI. 19 (fig. 2)

- incomplete specimen

BSPG AS I 974 - Oppel (1862), PI. 19 (fig. 3)

- length of thè specimen: 4 cm

BSPG AS I 976 - Oppel (1862), PI. 19 (fig. 4)

- length of thè specimen: 3 cm

BSPG AS VII 1429 - Oppel (1862), PI. 19 (fig. 1)

- length of thè specimen: 5 cm

BSPG AS I 975 - Oppel (1862), PI. 19 (fig. 5)

- length of thè specimen: 2.5 cm

BSPG AS VII 732 - Oppel (1862), PI. 14 (fig. 5)

- length of thè specimen: 2 cm

BSPG AS VII 733 - Oppel (1862), PI. 14 (fig. 6)

- length of thè specimen: 1 cm

BSPG AS VII 731 - Oppel (1862), PI. 14 (fig. 3)

- length of thè specimen: 4.5 cm

Description. Median-sized glypheid with strongly

tuberculate exoskeleton.

Carapace. Carapace cylindrical laterally flattened and

strongly tuberculate. Rostrum very short and edentate

with curved distai extremity. Cervical and branchiocar-

diac grooves deep. Postcervical groove weak, joined to

branchiocardiac groove near their lower ends. Antennal,

hepatic, and ventral grooves weak. Antennal spine well

developed. Three/four longitudinal carinae strongly

tuberculate extend parallel in gastric and antennal

regions. Ocular incision narrow with weakly developed

antennal and ptergostomial angles.

Abdomen. Somite I-V subrectangular of equal

h ngth and with tuberculate tergal surface. Somites I-V

w ith subtriangular pleurae. Telson subrectangular with

rounded distai extremity and three tuberculate carinae

extending its total length. Protopod subrectangular.

Uropodal endopod smooth. Uropodal exopod with weak

median longitudinal groove, marked by two rows of

spines, two different distai spines on thè outer margin

(one longer and one shorter), outer margin with small

tubercules, and rounded diaeresis with upper serrate

margin.

Cephalic appendages. Eyestalk very elongate and

strong. 3rd maxilliped very elongate with first article

having a strong distai spine on thè outer margin and

second article with two strong distai spines on thè

outer margin respectively. Antennular articles thin

and elongate of which thè distai one carries two multi-

articulate flagella of equal length. lst and 3rd antennal

articles short, 2nd antennal article very elongate with

dorsal surface covered with rows of strong parallel

spines. Multiarticulate flagellum elongate, twice thè

body length. Scaphocerite triangular with pointed distai

extremity and lower margin with a row of thin and elon¬

gate spines.

Thoracic appendages. Pereiopod I subchelate with

strong, elongate subrectangular merus and propodus

and short subtriangular carpus. Movable finger thin

and elongate with tuberculate dorsal surface and ser¬

rate upper and lower margins. Lower margin of pro¬

podus with strong spines of equal length in proximal

and mediai parts; distai part with two spines, first

stronger and more elongate than second. Dorsal surface

of merus, carpus, and propodus covered with strong

spines, locateci along margins in merus, and in parallel

rows in carpus and propodus. Pereiopods II-V achelate.

Lower margin of merus and carpus of pereiopods II-IV

with short spines.

Abdominal appendages. Pleopods with an unseg-

mented peduncle which carries two elongate multiarticu¬

late flagella.

Discussion. Feldmann & de Saint Laurent (2002)

gave a check list of 29 species included in Glyphea

v. Meyer, 1835. Among these species, G. squamosa

(Munster, 1839), is now considered thè type species of

Squamosoglyphea Beurlen, 1930 (see thè description

and discussion in Schweigert & Garassino, 2005, and

below). Moreover, thè authors did not consider seven

species, described by Damborenea & Mancenido (1987),

Garassino (1996, 1997, 2000, 2001), Polz (2000), and

Schweitzer & Feldmann (2001): G. eureka Damborenea

& Mancenido, 1987, G. tricarinata Garassino, 1996, G.

tonelloi Garassino, 1997, G. rigoi Garassino, 2000, G.

viohli Polz, 2000, G. damesi Garassino, 2001, and G.

michelae Schweitzer & Feldmann, 2001.

As reported by Glaessner (1929), Glyphea may include

more species. However, since this genus has never been

reviewed, probably many species could be synonym for

some morphological characters, such as thè path of thè

grooves, thè number of carinae in gastric and antennal

regions and thè structure of pereiopod I. Unfortunately,

this review is made difficult for thè problems of finding

thè originai samples and for thè loss of many originai

specimens.

On thè basis of this reflection, we consider Feld¬

mann & de Saint Laurent’s paper as thè most updated

paper known to date. So today, Glyphea includes 36

species from thè Upper Triassic (Norian) to Eocene

(Bartonian) of Europe (Germany, France, England,

Poland, ltaly and Spain), Lebanon, East Africa, North

and South America, New Zealand, Australia, and Ant-

arctic Peninsula.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART 1

17

Glyphea tenuis Oppel, 1861

Fig. 9, PI. IV (fig. 1), PI. XIII (fig. 4)

1861 - Glyphea tenuis Oppel; p. 110

1 862 - Glyphea tenuis Oppel; p. 76, PI. 20 (figs. 2-3)

1883 - Glyphea tenuis Oppel in Winckler; p. 27, Fig. 5

1904 - Glyphea tenuis Oppel in Walther; p. 174

1994 - Glyphea pseudoscyllarus (Schlotheim) in Frickhinger; p. 120,

Fig. 207

1994 - Glyphea tenuis Oppel in Frickhinger; p. 120, Fig. 208

2005b - Glyphea tenuis Oppel in Schweigert & Garassino; p. 498

Diagnosis: carapace cylindrical laterally flattened;

rostrum short and edentate with curved distai extremity;

cervical and branchiocardiac grooves deep; postcervi-

cal groove weak; antennal, hepatic, and ventral grooves

weak; one/two longitudinal carinae weak extend parallel

in gastric and antennal regions; pereiopod I subchelate;

pereiopods II-V achelate; uropodal exopod with diaer-

esis.

Holotype by monotypy: BSPG AS Vili 95 (Oppel,

1862 - PI. 20, figs. 2-3) housed in thè Bayerische

Staatssammlung fiir Palàontologie und Geologie in

Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Eichstàtt.

Material: Oppel (1862) reported one specimen,

ascribed to this species, and housed in thè Bayerische

Staatssammlung fiir Palàontologie und Geologie in

Munich where, today, it is stili preserved.

BSPG AS Vili 95 - Oppel (1862), PI. 20 (figs. 2-3)

- length of thè specimen: 4 cm

One specimen (n. cat. 9911 - Wulf collection) was

used to describe thè tail fan, poorly preserved in thè origi¬

nai specimen.

Description. Median-sized glypheid with finely tuber-

culate exoskeleton.

Carapace. Carapace cylindrical laterally flattened

and finely tuberculate. Rostrum short and edentate with

curved distai extremity. Cervical and branchiocardiac

grooves deep. Postcervical groove weak. Antennal,

hepatic, and ventral grooves weak. One/two longitudinal

carinae extend parallel in gastric and antennal regions.

Ocular incision narrow with weakly developed antenna!

and ptergostomial angles.

Abdomen. Somites I-V subrectangular of equal

length and with smooth tergal surface. Pleurae of

somites I-V ending with a point in thè median part.

Telson subrectangular with rounded distai extremity.

Protopod subrectangular. Uropodal endopod with weak

longitudinal carina. Uropodal exopod with weak longi¬

tudinal carina, strong distai spine in thè outer margin,

and rounded diaeresis.

Cephalic appendages. Eyestalk elongate and strong.

3rd maxilliped not preserved. Antennular articles thin

and elongate of which thè distai one carries two multi-

articulate flagella of equal length. lst and 3rd antennal

articles short and 2nd antennal article very elongate.

Multiarticulate flagellum elongate, twice thè body

length. Scaphocerite triangular with pointed distai

extremity.

Thoracic appendages. Pereiopod I subchelate with

short subrectangular merus and propodus and short sub-

triangular carpus. Movable fìnger thin and elongate with

smooth dorsal surface. Lower margin of propodus with

two/three strong spines of equal length distally. Dorsal

surface of merus, carpus, and propodus tuberculate.

Pereiopods II-V achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. Oppel (1862) pointed out thè main

morphological differences of this species respect to G.

pseudoscyllarus (Schlotheim, 1822) in order to justify

its institution: smooth exoskeleton, shorter antennal

articles and shorter eyestalk. The study of thè originai

specimen confirmed thè systematic validity of G. tenuis

Oppel, 1860: postcervical and branchiocardiac grooves

extend parallel without joining near their lower ends,

as in G. pseudoscyllarus ; one/two weak longitudinal

carinae in gastric and antennal regions unlike G. pseu¬

doscyllarus in which three/four tuberculate carinae are

present in these regions; different shape of pereiopod

I with shorter merus and propodus and dorsal surface

of carpus and propodus without strong spines arranged

in parallel rows, as in G. pseudoscyllarus ; merus and

carpus of pereiopods II-IV without spines along thè

outer margins, well developed instead in G. pseudoscyl¬

larus.

Genus Squamosoglyphea Beurlen, 1930

Type species: Glyphea dressieri v. Meyer, 1840 (for

correct spelling of this taxon see Schweigert & Garassino,

2005a)

Squamosoglyphea squamosa (Mtinster, 1839)

Fig. 9, PI. IV (fig. 2), PI. XIII (fig. 5)

1839 - Orphnea squamosa Mtinster; p. 41, PI. 14 (fig. 4)

1 853 - Orphnea squamosa Mtinster in Frischmann; p. 29

1 854 - Orphnea squamosa Mtinster in Pictet; p. 449, PI. 42 (fig. 9)

1861 - Glyphea squamosa (Mtinster) in Oppel; p. 110

1862 - Glyphea squamosa (Mtinster) in Oppel; p. 75, PI. 20 (fig. 1)

1904 - Glyphea squamosa (Mtinster) in Walther; p. 174

1925 - Glyphea squamosa (Mtinster) in Van Straelen; p. 189

1928 - Glyphea squamosa (Mtinster) in Beurlen; p. 136

1929 - Glyphea squamosa (Mtinster) in Glaessner; p. 191

1961 - Glyphea squamosa (Mtinster) in Martin; p. 70

1973 - Glyphea squamosa (Mtinster) in Forster; p. 34

2001 - Glyphea squamosa (Mtinster) in Schweitzer & Feldmann; p.

177

2005a - Squamosoglyphea squamosa (Mtinster) in Schweigert & Ga¬

rassino; p. 274, Figs. 1-2

The poor state of preservation of thè only originai

specimen, ascribed to this species, makes it difficult to

extend its morphological description over thè originai one

by Mtinster.

The possibility to clean thè tail fan allowed description

of it for thè first time. Some more details became obvious

from a topotype.

Diagnosis: carapace cylindrical laterally flattened

having wide superimposed scales; rostrum short and

edentate; cervical, postcervical, and branchiocardiac

grooves weak; pereiopod 1 subchelate with propodus

longer than wide, spineless on thè lower margin; uropodal

exopod with diaeresis.

18

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

Holotype by monotypy: BSPG AS VII 734 (Miinster,

1839 - PI. 14, fig. 4), housed in thè Bayerische Staatssa-

mmlung fìir Palàontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian).

Type locality: Kelheim (after lithology; originally

labelled “Solnhofen”).

Material: Oppel (1862) reported two specimens

belonging to this species, so divided: one from thè Reden-

bacher collection and one (thè originai of Miinster) from

thè collection of thè Bayerische Staatssammlung fur Palà¬

ontologie und Geologie in Munich. We studied thè holo¬

type by Miinster, illustrated by Miinster (1839) and Oppel

(1862), and thè other specimen from thè Redenbacher

collection in thè Museum fur Naturkunde der Humboldt-

Universitàt in Berlin.

BSPG AS VII 734 - Miinster (1839), PI. 14 (fìg. 4)

BSPG AS VII 734 - Oppel (1862), PI. 20 (fìg. 1)

- length of thè specimen: 5 cm

MNHB MB. A. 1 1 06 - Oppel ( 1 862), p. 75, not figured

- length of thè specimen: 4.2 cm

Description. Small-sized glypheid with exoskeleton

characterized by imbricated scales. The holotype is dor-

sally embedded, whereas thè second specimen is embed-

ded in oblique view.

Carapace. Carapace cylindrical laterally flattened

having wide superimposed scales in cardiac and branchial

regions reducing in size toward gastric and antennal

regions. Rostrum short and edentate. Cervical, postcervi-

cal, and branchiocardiac grooves weak. Ocular incision

narrow with weakly developed antennal and ptergosto-

mial angles. Posterior margin sinuous with a thin mar¬

ginai carina.

Abdomen. Somites I-V subrectangular of equal

length and with smooth tergal surface. Pleurae of

somites I-V ending with a point in thè median part.

Somite VI subrectangular longer than thè previous

ones. Telson subrectangular with rounded distai extrem-

ity. Dorsal surface of telson with a pair of narrow and

raised carinae originating in proximal part of lateral

margins and converging in thè middle without joining.

Rounded basai tubercle in thè middle between thè two

carinae. A pair of grooves narrow and weak arise from

thè distai end of thè carinae diverging toward thè distai

extremity. Protopod subrectangular. Uropodal endopod

with weak longitudinal groove marked with two rows of

rounded tubercles. Uropodal exopod with deep longitu¬

dinal groove marked with two rows of rounded tuber¬

cles, strong spine in thè distai part of outer margin, and

subrounded diaeresis. In thè ventral part of thè tergum

ol thè somites, forming forwardly directed lobes are

developed, located just above thè pleurae, like in thè

reconstruction of S. udressieri by Oppel (1862 - PI

16, fìg. 7).

Cephalic appendages. Merocerite of antennae strongly

elongate with tour thin and elongate spines along lower

margin. Carpocerite of antennae short with elongate

multiarticulate lìagellum (total length unknown). Anten¬

nule poorly preserved, both flagella of different length

(ca. 5 mm vs. 6 mm).

Thoracic appendages. Pereiopod I subchelate with

piopodus longer than wide and spineless lower margin.

Movable finger thin and elongate with a median groove.

Carpus short and subtriangular with spineless lower

margin. Surface of propodus with irregularly arranged

scale-like rounded tubercles with forwardly directing

tips. Surface of carpus with scale-like tubercles exhibiting

multiple forwardly directing tips, especially in thè distai

part. Pereiopods II-V achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. Beurlen (1930) based a new subgenus on

thè closely related S. dressieri (v. Meyer, 1840) - for thè

correct spelling of thè latter see Schweigert & Garassino

(2005a). The presence of superimposed scales instead

of tubercles is so different from Glyphea sensu strido

that we give genus rank to this taxon. Besides S. squa¬

mosa, two other species of this genus occur in thè Upper

Jurassic lithographic limestones of S Germany which

were described separately (Schweigert & Garassino,

2005).

Family Mecochiridae Van Straelen, 1924

Genus Mecochirus Germar, 1827

Type species: Macrourites Iongimanatus Schlotheim,

1820

Solnhofen species: Mecochirus Iongimanatus

(Schlotheim, 1820); Mecochirus dubius (Miinster,

1839)

Mecochirus Iongimanatus (Schlotheim, 1820) ($)

Fig. 9, PI. V (fìg. 1), PI. XIII (fìg. 6), PI. XIV (fìg. 1)

1820 - Macrourites Iongimanatus Schlotheim; p. 38

1 822 - Macrourites Iongimanatus Schlotheim; p. 33

1 822 - Macrourites Iongimanatus Schlotheim in Brogniart & Desmar-

est; p. 136, PI. 5 (fig. 10)

1 823 - Macrourites Iongimanatus Schlotheim; p. 56

1 825 - Palaemon Iongimanatus Kriiger; p. 130

1 827 - Mecochirus locusta Germar; p. 102

1827 - Mecochirus Bajeri Germar; p. 103, Text-fig. 5, nov. syn.

1 829 - Palaemon Iongimanatus Kriiger in Holl; p. 152

1837 - Megachirus locusta Bronn; p. 476, PI. 27 (fig. 1)

1837 - Megachirus longimanus Bronn; p. 476, Text-fig. 16a

1839 - Megachirus brevimanus Miinster; p. 34, PI. 13 (figs. 1?, 2, 3),

nov. syn. (c?)

1 839 - Megachirus locusta Bronn in Miinster; p. 3 1 , PI. 1 1 (figs. 1 -5)

1 839 - Megachirus Bajeri (Germar) in Miinster; p. 33, PI. 12 (figs. 1-5),

PI. 13 (figs. 6-7)

1839 - Megachirus fimbriatus Miinster; p. 35

1839 - Megachirus intermedius Miinster; p. 35, PI. 13 (figs. 4?, 5)

1839 - Pterochirus remimanus Bronn in Miinster; p. 27, PI. 16 (figs.

1-2)

1839 - Pterochirus elongatus Miinster; p. 28, PI. 16 (fig. 3)

1839 - Orphnea longimanus Miinster; p. 42, PI. 14 (fig. 7)

1 848 - Megachirus locusta Bronn; p. 708

1850 - Mecochirus locusta Germar in Quenstedt; p. 196, PI. 2 (figs.

1-3)

1853 - Mecochirus locusta Germar in Bronn; p. 418, PI. 27 (fig. 1,

16a)

1862 - Mecochirus longimanus (Schlotheim) in Oppel; p. 82, PI. 22

(fig. 4), PI. 23 (fig. 1)

1 862 - Mecochirus Bajeri Germar in Oppel; p. 83, PI. 23 (fig. 2)

1862 - Mecochirus brevimanus (Miinster) in Oppel; p. 84, PI. 22 (figs.

5-6)

1904 - Mecochirus longimanus (Schlotheim) in Walther; p. 174

THE UPPER JURASS1C SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

19

1904 - Mecochirus Bajeri Germar in Walther; p. 174

1904 - Mecochirus brevimanus (Miinster) in Walther; p. 174

1925 - Mecochirus longimanatus (Schlotheim) in Van Straelen; p. 218

1929 - Mecochirus longimanatus (Schlotheim) in Glaessner; p. 250

1929 - Mecochirus Bajeri Germar in Glaessner; p. 249

1929 - Mecochirus brevimanus (Miinster) in Glaessner; p. 249

1969 - Mecochirus longimanatus (Schlotheim) in Glaessner; R 464,

Text-fig. 270/1

1971 - Mecochirus longimanatus (Schlotheim) in Forster; p. 403,

Text-fig. 5

1994 - Mecochirus longimanatus (Schlotheim) in Frickhinger; p. 122,

Figs. 214-216

1994 - Mecochirus bajeri Germar in Frickhinger; p. 122, Fig. 212

1994 - Mecochirus brevimanus (Miinster) in Frickhinger; p. 122, Fig. 213

1996 - Mecochirus sp. in Ròper, Rothgaenger & Rothgaenger; p. 72,

Fig. 89

1999 - Mecochirus brevimanus (Miinster) in Frickhinger; p. 43, Fig. 68

2000 - Mecochirus brevimanus (Miinster) in Ròper, Rothgaenger &

Rothgaenger; p. 91, Fig. 105

2005b - Mecochirus longimanatus (Schlotheim) in Schweigert & Ga-

rassino; p. 498

Diagnosis: carapace cylindrical laterally flattened;

rostrum short and edentate; cervical groove deep in thè

third anterior of carapace; antennal and hepatic grooves

deep; postcervical and branchiocardiac grooves weak

extending parallel; three carinae strong and tuberculate

in thè gastric and antennal regions; extraordinary exten-

sion of pereiopod I subchelate; pereiopod II subchelate;

pereiopods III-V achelate; uropodal exopod with diaer-

esis.

Neotype (originai type series of Schlotheim lost):

Macrourites longimanatus (= Mecochirus longimanatus ),

BSPG AS V 45, housed in thè Bayerische Staatssammlung

fìir Palàontologie und Geologie in Munich.

Olir studies revealed that Mecochirus bajeri from

Solnhofen and figured by Germar (1827 - PI. 1, fig. 5)

is synonym with Mecochirus longimanatus , as discussed

below. The monotypic holotype was not traceable in thè

collection of thè Institut fur Geowissenschaften in Halle/

Saale (Germany) (Dr. N. Hauschke, Halle, pers. com.),

and it is probably lost.

Our studies revealed that Megachirus brevimanus

from Eichstàtt and figured by Miinster (1839 - PI. 13,

fig. 2) is synonym with Mecochirus longimanatus , as

discussed below. This specimen, BSPG ASVII 739, is

thè lectotype of Miinster’s species, housed in thè Bay¬

erische Staatssammlung fur Palàontologie und Geologie

in Munich.

Stratigraphic range; Upper Jurassic (upper Kim-

meridgian - lower Tithonian).

Type locality: Solnhofen.

Material: Schlotheim (1820) introduced thè name

reported on a sample of three specimens and com-

mented that this species is thè most common one in thè

Solnhofen shales. A neotype was selected because none

of thè specimens of Schlotheim collection was traceable

in thè Museum fur Naturkunde in Humboldt University

in Berlin. Oppel (1862) reported 103 specimens belong-

ing to this species, so divided: 30 from thè Redenbacher

collection and 73 from thè collection of thè Bayerische

Staatssammlung fur Palàontologie und Geologie in

Munich. W e studied of this originai sample six speci¬

mens, housed in thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich in which we

identified three specimens, illustrated by Miinster (1839)

and three specimens, illustrated by Oppel (1862). Two

specimens were identified as belonging to Mùnster’s

originai collection thanks to thè 19,h century labels. We

report thè list of thè specimens, studied by Miinster and

Oppel:

BSPG AS VII 745 - Miinster (1839), PI. 11 (fig. 3)

- length of thè specimen: 6 cm

BSPG AS VII 746 - Miinster (1839), PI. 11 (fig. 1)

- length of thè specimen: 9 cm

BSPG AS VII 748 - Miinster (1839), PI. 16 (fig. 3)

- length of thè specimen: 4 cm

BSPG AS V 45 - Oppel (1862), PI. 22 (fig. 4)

- length of thè specimen: 1 0 cm

BSPG AS I 780 - Oppel (1862), PI. 22 (fig. 5)

- length of thè specimen: 5 cm

BSPG AS VI 31 - Oppel (1862), PI. 23 (fig. 1)

- length of thè specimen: 8 cm

Oppel ( 1 862) reported 7 0 specimens which he assigned

to Mecochirus bajeri Germar, 1 827, so divided: 20 from

thè Redenbacher collection and 50 from thè collection of

thè Bayerische Staatssammlung frir Palàontologie und

Geologie in Munich. We studied of this originai sample

nine specimens, housed in thè Bayerische Staatssa¬

mmlung tur Palàontologie und Geologie in Munich

in which we identified eight specimens, illustrated by

Miinster (1839), and one specimen, illustrated by Oppel

(1862). We report thè list of thè specimens, studied by

Miinster and Oppel:

BSPG AS VII 735 - Miinster (1839), PI. 16 (fig. 2)

- length of thè specimen: 5 cm

BSPG AS VII 736 - Miinster (1839), PI. 16 (fig. 1)

- incomplete specimen

BSPG AS VII 738 - Miinster (1839), PI. 12 (fig. 5)

- length of thè specimen: 7 cm

BSPG AS VII 741 - Miinster (1839), PI. 12 (fig. 3)

- length of thè specimen: 6 cm

BSPG AS VII 742 - Miinster (1839), PI. 12 (fig. 4)

- incomplete specimen

BSPG AS VII 743 - Miinster (1839), PI. 13 (fig. 4)

- length of thè specimen: 6 cm

BSPG AS VII 744 - Miinster (1839), PI. 13 (fig. 5)

- length of thè specimen: 6 cm

BSPG AS VII 749 - Miinster (1839), PI. 13 (fig. 6)

- length of thè specimen: 6 cm

BSPG AS I 981 - Oppel (1862), PI. 23 (fig. 2)

- length of thè specimen: 8 cm

Oppel (1862) reported 50 specimens which he

assigned to Mecochirus brevimanus (Miinster, 1839)

without pointing out how many specimens belonged to

thè Redenbacher collection and to thè collection of thè

Bayerische Staatssammlung fur Palàontologie und Geolo¬

gie in Munich. We studied three specimens of this origi¬

nai sample, housed in thè Bayerische Staatssammlung fiir

Palàontologie und Geologie in Munich and illustrated by

Miinster (1839). One specimen was identified as belong¬

ing to Miinster ’s originai collection thanks to thè 191*1 cen¬

tury labels. We report thè list of thè specimens, studied by

Miinster:

BSPG AS VII 739 - Miinster (1839), PI. 13 (fig. 2)

- length of thè specimen: 7 cm

BSPG AS VII 740 - Miinster (1839), PI. 13 (fig. 1)

- length of thè specimen: 10 cm

20

ALESSANDRO GARASSINO & GONTER SCHWEIGERT

BSPG AS VII 750 - Miinster (1839), PI. 13 (fig. 3)

- length of thè specimen: 4 cm

Description. Large mecochirid with slightly tubercu-

late exoskeleton.

Carapace. Carapace cylindrical laterally flattened

having straight dorsal margin. Rostrum short and eden-

tate. Cervical groove deep in thè third anterior of cara¬

pace. Antennal and hepatic grooves deep. Postcervical

and branchiocardiac grooves weak and parallel. Three

carinae strong and tuberculate in gastric and antennal

regions. Ocular incision narrow, and shallow.

Abdomen. Somites I-VI subrectangular of equal

length with slightly tuberculate tergal surface. Pleurae

of somites I-V subtriangular with serrate margins. Telson

subrectangular with one raised tubercle in proximal part

of dorsal surface, a pair of thin spine in thè distai part of

lateral margins, a thin median spine on lower margin and

rounded distai extremity. Protopod subrectangular. Uro-

podal endopod with a strong median longitudinal carina.

Uropodal exopod with a strong median longitudinal

carina and a rounded diaeresis with upper serrate margin

having one longer median spine. Fringed lower margins

of uropodal endopod and exopod.

Cephalic appendages. Eyestalk short. Antennulae not

preserved. Antennal articles strong, and elongate of which

thè distai one carries a strong elongate multiarticulate

flagellum, twice thè body length. Scaphocerite triangular

with pointed distai extremity.

Thoracic appendages. Extraordinary extension ofperei-

opod I subchelate. Possibly this is a character of female

specimens (see M. brevimanus below supposed to be thè

corresponding male). Lower margin of propodus with one

strong and sharp distai tooth. Movable finger strong and

elongate with tuberculate upper and lower margins. Dorsal

surface of movable finger with two parallel rows of small

tubercles. Fringed lower margin of merus, carpus, and

propodus of pereiopod I. Fringed upper and lower margins

of movable finger of pereiopod I. Pereiopod II subchelate

with strong and stout propodus. Fringed lower margins of

merus, carpus, and propodus of pereiopod II. Pereiopods

III-V of equal length and achelate.

Abdominal appendages. Pleopods with an unseg-

mented peduncle which carries two elongate multiarticu¬

late flagella.

Discussion. Forster (1971) reviewed this genus,

including 18 species. Later, Simpson & Middleton (1985),

studying a sample of mecochirids from thè Lower Creta-

ceous of England, described thè new genus Meyerella

having as type species Mecochirus magnus McCoy, 1849.

Moreover, thè authors, using as reference Forster (1971),

pointed out that Mecochirus schwarzi (Kitchin, 1913)

is synonym with Mecochirus rapax (Harbort, 1905),

better referred to Meyerella , and Mecochirus mexicanus

(Rathbun, 1935) and Mecochirus bolivari (Van Strae-

len, 1927) are synonyms of Meyerella magna (McCoy,

1849). Finally, thè authors pointed out that Mecochirus

ornatus (Phillips, 1829) was thè type species of Meyeria

McCoy, 1849, and Mecochirus gracilis (Glaessner, 1932)

belongs to this genus. Today, Mecochirus Germar, 1827,

'• n from thè Lower Jurassic (Sinemurian) to thè

Upper Cretaceous (Maastrichtian) of Europe (Germany,

France, England and Italy), Greenland, South America

(Chile), Antarctic Peninsula and New Zealand, includes

fifteen species. Three species are known from thè Lower

Jurassic: M. olifex Quenstedt, 1856 (Germany - Sinemu¬

rian), M. germari Garassino, 1996 (Italy - Sinemurian),

and M. eckerti Frentzen, 1937 (Germany - Toarcian).

Three species have been recognized from thè Middle

Jurassic: M. clypeatus (Carter, 1898) (England - Batho-

nian), M. socialis (v. Meyer, 1841) (Germany, France,

England - Callovian), and M. pearcei McCoy, 1849

(England - Oxfordian). Seven species are described from

thè Upper Jurassic: M. peytoni Woodward, 1876 (France,

England - Kimmeridgian), M. leionotus (Krause, 1891)

(Germany - Kimmeridgian), M. minimus (Hée, 1924)

(France - Kimmeridgian), M. marwicki Glaessner, 1960

(New Zealand - Kimmeridgian), M. chilensis Forster &

Hillebrandt, 1984 (Chile - Kimmeridgian), and M. longi-

manatus (Schlotheim, 1822) (Germany - Tithonian). One

is known from thè Lower Cretaceous: M. houdardi Van

Straelen, 1936 (France - Albian), and two are reported

from thè Upper Cretaceous: M. crofti Ball, 1960 (Antarc¬

tic Peninsula - Campanian), and M. rostratus Collins &

Rasmussen, 1 992 (Greenland - Maastrichtian).

Germar (1827), describing Mecochirus bajeri on one

incomplete specimen, pointed out thè main morphologi-

cal differences respect M. longimanatus, such as a shorter

pereiopod I and thè abdomen longer than carapace. Oppel

( 1 862) pointed out that thè shorter pereiopod I is thè main

morphological characters distinguished M. bajeri from

thè type species. However, thè study of specimens of M.

longimanatus and M. bajeri, having thè same dimensionai

class, showed that thè length of pereiopod I was thè same,

as well as thè length of thè abdomen. Moreover, thè speci¬

mens belonging to M. bajeri showed thè same morpholog¬

ical characters of thè type species, such as thè tuberculate

longitudinal carinae of carapace, a strong distai spine of

thè propodus of pereiopod I and thè same structure of tail

fan, enough to consider it as synonym of M. longimanatus,

probably a juvenile stage of thè type species.

Miinster (1839), describing Megachirus brevimanus,

pointed out thè main morphological differences respect

M. longimanatus, such as thè shorter pereiopod I and pro¬

podus of pereiopod I shorter than carapace. Apart from

these very significai characters, thè specimens belonging

to M. brevimanus exhibit thè same morhological charac¬

ters of thè type species, such as thè tuberculate longitudi¬

nal carinae of carapace, thè strong distai spine of thè pro¬

podus of pereiopod I, and thè same structure of tail fan.

After these remarkable characters in common between

M. longimanatus and M. brevimanus, we suppose these

forms represent sexual dimorphs of a single species. In

fact, in Recent glypheid Neoglyphea inopinata, thè clos-

est living relative of this group of decapods, thè sexual

dimorphism also affects thè lengths of pereiopod I and its

articles, with thè longer ones are developed in thè female

(Forest & de Saint Laurent, 1981, 1989). Thus we sup¬

pose M. brevimanus is thè male and M. longimanatus thè

female form.

Mecochirus dubius (Miinster, 1839)

PI. XIV (fig. 2)

1839 - Pterochirus dubius Miinster; p. 29, PI. 16 (figs. 4-6)

1 840 - Noma lithophila Miinster; p. 22, PI. 3 (fig. 9)

1862 - Mecochirus dubius (Miinster) in Oppel; p. 84, PI. 23 (fig. 3)

1929 - Mecochirus dubius (Miinster) in Glaessner; p. 249-250

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

21

Lectotype: BSPG AS I 979 (Oppel, 1 862 - PI. 23, fig.

3), housed in thè Bayerische Staatssammlung fìir Palàon-

tologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported 11 specimens

belonging to this species from thè collection of thè Bay¬

erische Staatssammlung fìir Palaontologie und Geologie

in Munich. We studied of this originai sample three speci¬

mens in which we identified two specimens, illustrated by

Miinster (1839) and one specimen, illustrated by Oppel

(1862). Four specimens were identified as belonging to

Miinster ’s originai collection thanks to thè 19th century

labels. We report thè list of thè specimens, studied by

Miinster and Oppel:

BSPG AS VII 737 - Miinster (1839), PI. 16 (fig. 6)

- length of thè specimen: 2 cm

BSPG AS VII 747 - Miinster (1839), PI. 16 (fig. 4)

- length of thè specimen: 2 cm

BSPG AS I 979 - Oppel (1862), PI. 23 (fig. 3)

- incomplete specimen

Discussion. Miinster (1839) justified thè institution

of this species because pereiopod I is shorter and higher

than that of thè other species, pointing out thè difficulty of

giving its morphological description for thè fragmentary

of thè sample. The study of thè originai specimens stili

available has pointed out that their poor state of preserva-

tion and their ventral view made difficult to give not only

a morphological description of M. dubius, but also a com-

parison with thè other species belonging to Mecochirus.

Even though thè small sizes of thè specimens lead us to

think that they could be ontogenetic stages of M. longi-

manatus, it is difficult to assert it.

Infraorder Thalassinidea Latreille, 1831

Superfamily Axioidea Huxley, 1879

Family Axiidae Huxley, 1879

Genus Etallonia Oppel, 1861

Type species: Maglia longimana Miinster, 1839

Etallonia longimana (Miinster, 1839)

Figs. 3, 9, PI. XIV (fig. 3)

1839 -Magila longimana Miinster; p. 25, PI. 10 (fig. 3)

1861 - Etallonia longimana (Miinster) in Oppel; p. 361

1862 - Etallonia longimana (Miinster) in Oppel; p. 49, PI. 12 (figs.

5-7)

1904 - Etallonia longimana (Miinster) in Walther; p. 175

1925 - Etallonia longimana (Miinster) in Van Straelen; p. 231, Text-

fig. 109

1929 - Etallonia longimana (Miinster) in Glaessner; p. 168

1930 - Etallonia longimana (Miinster) in Beurlen & Glaessner; p. 78

1969 - Etallonia longimana (Miinster) in Glaessner; R 477, Text-fig.

282/4

1994 - Etallonia longimana (Miinster) in Frickhinger; p. 120, Fig. 205

1999 - Etallonia longimana (Miinster) in Polz; p. 34, Text-fig. 2, PI. 1

(figs. 5-7)

2005b - Etallonia longimana (Miinster) in Schweigert & Garassino; p. 499

The poor state of preservation of thè originai speci¬

mens and thè lack of better preserved new specimens did

not permit adding new morphological characters to thè

originai description by Miinster. In fact, thè carapace,

cephalic appendages, and tail fan are too poorly preserved

to give their description (Fig. 3).

Dignosis: carapace cylindrical laterally flattened; perei¬

opod I subchelate; pereiopods II with small chela; pereio-

pods III-V achelate; uropodal exopod with diaeresis.

Lectotype: BSPG AS VII 259 (Miinster, 1839 - PI.

10, fig. 3), housed in thè Bayerische Staatssammlung fìir

Palaontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported five specimens

belonging to this species, so divided: two from thè

Redenbacher collection and three from thè collection of

thè Bayerische Staatssammlung fìir Palaontologie und

Geologie in Munich. We studied of this originai sample

two specimens, housed in thè Bayerische Staatssammlung

fìir Palaontologie und Geologie in Munich in which we

identified one specimen, illustrated by Miinster (1839)

and one specimen, illustrated by Oppel (1862). We report

thè list of thè specimens, studied by Miinster and Oppel:

BSPG AS VII 259 - Miinster (1839), PI. 10 (fig. 3)

BSPG AS VII 259 - Oppel (1862), PI. 12 (fig. 5)

- length of thè specimen: 3 cm

BSPG AS Vili 96 - Oppel (1862), PI. 12 (fig. 7)

- length of thè specimen: 2 cm

Description. Small-sized axiid with smooth exoskel-

eton.

Carapace. Carapace cylindrical laterally flattened.

Poorly preserved in both specimens.

Abdomen. Somites I-V subrectangular of equal length

with rounded pleurae. Somite VI subrectangular longer

than thè previuos ones. Telson subrectangular with

rounded distai extremity. Uropodal exopod with weak

longitudinal median carina and rounded diaeresis.

Cephalic appendages. Antennal flagellum elongate.

Thoracic appendages. Pereiopod I stronger than

thè others and subchelate. Movable finger pointed and

curved downward. Lateral spine (= fixed finger) straight

and shorter than movable finger. Basai spine in more or

less median position, forming with sinus fixed fìnger.

Propodus of pereiopod I with strong longitudinal median

carina. Pereiopods II elongate with small chela. Pereio¬

pods III-V elongate and achelate.

Abdominal appendages. Not preserved.

Discussion. Etallonia, very rare in thè lithographic

limestone, was reviewed by Forster (1977). This genus,

known only in thè Upper Jurassic (Oxfordian - Titho¬

nian) of Germany, Poland, and Great Britain, includes

- after Forster (1977) - four species, E. subtilis (Krause,

1908) (Poland - Oxfordian), E. isochela (Woodward,

1876) (Great Britain - Kimmeridgian), E. longimana

and E. hoellorum Polz, 1999 (Germany - Kimmeridgian/

Tithonian).

Forster (1977) synonymized Etallonia and Protaxius.

Maybe he was in error when he made that. The shape of

thè chelae of Callianassa isochela Woodward, 1876 (type

species of Protaxius Beurlen, 1930) is dose to that of P.

suevicus Beurlen, 1930. Forster (1965) took P. suevicus as

synonym of E. suprajurensis (Quenstedt, 1 852). We point

out that probably this is not correct because P. suevicus

lacks thè spine on thè subchela, typical of Etallonia. So

Fòrster’s results have to be reviewed. Two indeterminate

22

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

Fig. 3 - Etallonia longimana (Miinster, 1839), line drawing (F. Fogliazza).

species are known from thè Kimmeridgian and Tithonian

of Great Britain and Tanzania (Arkell, 1933; Beurlen,

1933). An incompletely known species from thè Toarcian

of Italy tentatively included in Etallonia by Garassino &

Teruzzi (2001) probably represents a Megachela Sch-

weigert, 2003.

Genus Magila Miinster, 1839

Type species: Magila latimana Miinster, 1839

Solnhofen species: Magila latimana Miinster, 1839;

Magila desmarestii (Miinster, 1839); ? Magila denticulata

Miinster, 1839

Magila latimana Miinster, 1839

Figs. 4, 9, PI. V (fig. 2), PI. XIV (figs. 4, 5)

1839 - Magila latimana Miinster; p. 25, PI. 10 (fig. 2)

1852 - Pagurus suprajurensis Quenstedt; p. 265, PI. 20 (fig. 9) (not

fig. 8)

1 858 - Pagurus suprajurensis Quenstedt; p. 804, PI. 99 (fig. 20)

1861- Magila latimana Miinster in Oppel; p. 361

1862 - Magila latimana Miinster in Oppel; p. 48, PI. 12 (figs. 1-2)

1862 - Magila suprajurensis (Quenstedt) in Oppel; p. 47

1885 - Pagurus suprajurensis Quenstedt; p. 405, PI. 31 (figs. 37-39)

(not fig. 36)

1891 - Callianassa suprajurensis (Quenstedt) in Krause; p. 209, PI.

14 (fig. 4)

1925 - Magila latimana Miinster in Van Straelen; p. 297

1928 - Magila latimana Miinster in Beurlen; p. 185

1928 Magila suprajurensis (Quenstedt) var. lata Beurlen; p. 185

1929 - Callianassa suprajurensis (Quenstedt) var. lata Beurlen in

Glaessner; p. 92

1930 - Magila suprajurensis (Quenstedt) in Beurlen; p. 229, Text-fig. 5

1965 - Magila latimana Miinster in Fòrster; p. 146, Text-fig. 2, PI. 2

(fig- 7)

1969 - Magila latimana Miinster in Glaessner; R 477, Text-fig. 282/3

1994 - Magila latimana Miinster in Frickhinger; p. 121, Fig. 211

2005b - Magila latimana Miinster in Schweigert & Garassino; p. 499

Diagnosis: Carapace cylindrical laterally flattened;

deep cervical groove strongly directed forward; one or

two carinae weak in antennal region; rostrum short and

edentate; antennal spine well developed; pereiopods I-III

chelate; pereiopod I larger and stronger than pereiopods

II-III; pereiopods IV-V achelate; uropodal exopod with

diaeresis.

Lectotype: BSPG AS VI 26 (Oppel, 1 862 - PI. 1 2, fig.

2), housed in thè Bayerische Staatssammlung ftir Palàon-

tologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: originally labelled as coming from

“Solnhofen”. According to lithology of thè rock matrix

containing thè lectotype it carne from thè vicinity of

Eichstàtt.

Material: Oppel (1862) reported ten specimens

belonging to this species, so divided: six from thè

Redenbacher collection and four from thè collection of

thè Bayerische Staatssammlung tur Palàontologie und

Geologie in Munich. We studied of this originai sample

four specimens, housed in Bayerische Staatssammlung

fiir Palàontologie und Geologie in Munich in which we

identified two specimens, illustrated by Miinster (1839)

and two specimens, illustrated by Oppel (1862). We

report thè list of thè specimens, studied by Miinster and

Oppel:

BSPG AS VII 199 - Miinster (1839), PI. 10 (fig. 5)

- length of thè specimen: 2.5 cm

BSPG AS VII 320 - Miinster (1839), PI. 10 (fig. 2)

- length of thè specimen: 2.5 cm

BSPG AS VI 26 - Oppel (1862), PI. 12 (fig. 2)

length of thè specimen: 2.5 cm

BSPG AS I 674 - Oppel (1862), PI. 12 (fig. lb)

- incomplete specimen

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

23

One specimen (n. cat. 0117 - Wulf collection) was

used to describe thè carapace and thè tail fan, poorly pre-

served in thè originai specimens.

Description. Small-sized axiid with weakly tubercu-

late exoskeleton.

Carapace. Carapace cylindrical laterally flattened.

Rostrum short, pointed, and edentate. One or two carinae

weak and straight in antennal region. Cervical groove

deep and median strongly directed forwards, reduc-

ing gastric and antennal regions. Cardiac and branchial

regions wide. Antennal spine well developed. Posterior

margin strengthened by a thin marginai carina.

Abdomen. Somites I-V subrectangular of equal length

with rounded pleurae. Somite VI longer than thè previ-

ous ones. Telson subtriangular, as long as uropods, with

pointed distai extremity. Uropodal endopod with a weak

median longitudinal carina ending with a strong spine on

lower margin and a strong distai spine on outer margin.

Uropodal exopod with a weak median longitudinal carina

ending with a small spine on lower margin and a strong

distai spine on outer margin. These spines marked a very

small diaeresis present only on outer half of lower margin.

Outer margin serrate of uropodal exopod (Fig. 4).

Fig. 4 - Magila latimana Miinster, 1839, tail fan, line drawing (F.

Fogliazza).

Cephalic appendages. Poorly preserved. Antennula

with two elongate and thin flagella. Antenna with a very

elongate and thick flagellum twice length of body.

Thoracic appendages. 3rd maxilliped not preserved.

Pereiopod I with stocky and short chelae with movable

and fixed fingers of equal length and with curved distai

extremity. Inner margins of movable and fixed fingers.

Pereiopods II-III with small and thin chelae. Pereiopods

IV-V elongate and achelate. Lower margin of propodus

of pereiopods IV-V with a row of thin spines thè last ones

stronger than thè previous ones.

Abdominal appendages. Pleopods with an unseg-

mented peduncle which carries two elongate multiarticu-

late flagella.

Discussion. Magila , very rare in thè lithographic

limestone, was reviewed by Forster (1977). This genus,

known from thè Toarcian to Tithonian (Lower - Upper

Jurassic) of Germany, France and Great Britain, includes

six species: M. bonjouri (Etallon, 1861) (France - Toar¬

cian), M. pichleri Oppel, 1862 (Germany, Great Britain

- Callovian), M. straeleni (Patrulius, 1959) (Romania,

Israel - Tithonian), M. latimana Miinster, 1839, M. des-

marestii (Miinster, 1839), and ?M denticulata Miinster,

1839 (Germany - Tithonian). Moreover, Forster (1977)

pointed out that M. prisca (Krause, 1891) must be con-

sidered a synonym of Callianassa krausei Bòhm, 1911.

Finally, Magila includes two subspecies, M. dura bi-

cristata Forster, 1977 (Poland - Oxfordian) and M. dura

dura (Moericke, 1889) (Russia - Tithonian).

Magila desmarestii (Miinster, 1839)

PI. XIV (fig. 6), PI. XV (fig. 1)

1839 -Aura desmaresti Miinster; p. 26, PI. 10 (fig. 5)

1854 -Aura desmarestii Miinster in Pictet; p. 454, PI. 42 (fig. 21)

1862 - Magila robusta Oppel; p. 48, PI. 12 (fig. 3), nov. syn.

1925 - Magila robusta Oppel in Van Straelen; p. 296

1929 - Magila robusta Oppel in Glaessner; p. 246

2000 - Magila cfr. atimana Miinster in Ròper, Rothgaenger & Roth-

gaenger; p. 91, Fig. 106

Holotype by monotypy: BSPG AS VII 199 (Miinster,

1839 - PI. 10, fig. 5), housed in thè Bayerische Staatssa-

mmlung fur Palàontologie und Geologie in Munich.

Our studies revealed that Magila robusta from Eich-

stàtt and figured by Oppel (1862 - PI. 12, fig. 3) is syno¬

nym with Magila desmarestii, as discussed below. This

specimen, AS I 675, is thè holotype of Oppel’s species,

housed in thè Bayerische Staatssammlung fur Palaontolo-

gie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported one specimen,

ascribed to this species, and housed in Bayerische

Staatssammlung fur Palàontologie und Geologie in

Munich where, today, it is stili preserved.

BSPG AS VII 199 - Miinster (1839), PI. 10 (fig. 5)

- length of thè specimen: 3.5 cm

Oppel (1862) reported thè presence of only one speci¬

men which he assigned to Magila robusta Oppel, 1862,

housed in thè Bayerische Staatssammlung fìir Palàon¬

tologie und Geologie in Munich where, today, it is stili

preserved.

BSPG AS I 675 - Oppel (1862), PI. 12 (fig. 3)

- incomplete specimen

Discussion. Miinster (1839) described this species

on one incomplete, small specimen. Even though, it is

ascribed to Magila for thè typical shape of thè chela of

pereiopod I, its poor state of preservation did not allow

a morphological description of M. desmarestii and thè

lack of diagnostic characters, typical of this species,

hardly allow comparison with thè other species of thè

genus.

Oppel (1862) described Magila robusta based on one

incomplete, small specimen. Even though, it is ascribed

to Magila for thè typical shape of thè chela of pereiopod

I, its poor state of preservation did not allow writing a

24

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

complete morphological description of M. robusta. How-

ever, thè shape of thè chelae of pereiopod I fits well with

those of M. desmarestii in order to suppose both species

as synonym.

IMagila denticulata Miinster, 1839

1839 -Maglia denticu/ata Miinster; p. 26, PI. 10 (fig. 4)

1862 - Magila denticu/ata Miinster in Oppel; p. 46

1925 - Magila denticu/ata Miinster in Van Straelen; p. 298

1929 - Eryma sp. in Glaessner; p. 160

Discussion. Miinster (1839) described this species

based on one incomplete, small specimen from Eichstàtt.

The loss of this specimen, probably housed in thè Bay-

erische Staatssammlung fur Palàontologie und Geologie

in Munich, made difficult to give a new morphological

description of Magila denticulata Miinster, 1839, in order

to establish its systematic validity, and to compare it with

thè other species of thè genus. So we consider Magila

denticulata a nomen dubium, not being able to establish

its validity or its probable synonymy.

As Van Straelen (1925) and Glaessner (1929) pointed

out, this species should be ascribed to Eryma Meyer,

1 840, for thè shape of chelae that resembles those of this

genus. The drawing by Miinster, however, hardly allows

such interpretation.

Indeterminate Family

Genus Orhomalus Etallon, 1861

Type species: Orhomalus virgulinus Etallon, 1861

Orhomalus deformis (Oppel, 1 862)

PI. XV (fig. 2)

1862 - Magila deformis Oppel; p. 49, PI. 12 (fig. 4)

1 925 - Orhomalus deformis (Oppel) in Van Straelen; p. 328

1 929 - Magila deformis Oppel in Glaessner; p. 245

1929 - Orhomalus deformis (Oppel) in Glaessner; p. 281

Holotype by monotypy: BSPG AS Vili 115 (Oppel,

1862 - PI. 12, fig. 4), housed in thè Bayerische Staatssa¬

mmlung fìir Palàontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported one specimen,

ascribed to this species, and housed in thè Bayerische

Staatssammlung tur Palàontologie und Geologie in

Munich where, today, it is stili preserved.

BSPG AS Vili 115 - Oppel (1862), PI. 12 (fìg.4)

- incomplete specimen

Discussion. Oppel (1862) described this species on

one incomplete specimen, ascribing it to Magila Mun-

ster, 1839, based upon thè morphological characters of

thè chelae. However, Van Straelen (1925) pointed out

that thè shape ot thè chelae of pereiopod I is typical of

Orhomalus Etallon, 1861. Even though thè studied speci¬

men is incomplete, it represents one of thè most complete

specimens of this genus since most species are based on

isolate chelae.

Infraorder Palinura Latreille, 1 803

Family Eryonidae De Haan, 1841

Genus Cycleryon Glaessner, 1 965

Type species: Macrourites propinquus Schlotheim,

1822

Solnhofen species: Cycleryon propinquus (Schlotheim,

1 822); Cycleryon orbicu/atus (Miinster, 1 839); Cycleryon

e/ongatus (Miinster, 1839); Cycleryon wulfi Garassino &

Schweigert, 2004

Cycleryon propinquus (Schlotheim, 1822) (c?)

Figs. 5, 6, 10, PI. VI, PI. XV (figs. 3, 4, 5, 6)

1822 - Macrourites propinquus Schlotheim; p. 35, PI. 3 (fig. 2)

1825 -Eryon Schlotheimi Kònig; p. 95, PI. 8 (fig. 93)

1 827 - Eryon propinquus (Schlotheim) in Germar; p. 99

1827 - Eryon spinimanus Germar; p. 99, nov. syn. ($)

1829 - Eryon Schlotheimi Kònig in Holl; p. 150

1836 -Eryon Schuberti v. Meyer; p. 271, PI. 12 (figs. 3, 6), nov. syn.

1836 - Eryon Schlotheimi Kònig in v. Meyer; p. 280

1838 - Eryon Rehmanni v. Meyer; p. 415

1 836 - Etyon speciosus Mùnster; p. 5, PI. 2, PI. 3 (fig. 2)

1 839 - Eryon Meyeri Miinster; p. 6, PI. 3 (fig. 1), PI. 4

1 839 - Eryon Schuberti v. Meyer in Miinster; p. 13, PI. 7 (figs. 6-9)

1839 - Eryon elongatus Mùnster; p. 9, PI. 5 (fig. 7)

1 852 - Eryon propinquus (Schlotheim) in Quenstedt; p. 267, PI. 20

(fig. 19)

1853 - Eryon propinquus (Schlotheim) in Frischmann; p. 26

1 853 - Eryon speciosus Mùnster in Frischmann; p. 26

1853 - Eryon Schuberti v. Meyer in Frischmann; p. 27

1855 - Eryon propinquus (Schlotheim) in Fraas; p. 92

1855 - Eryon spinimanus Germar in Fraas; p. 93

1 856 - Eryon spinimanus Germar in v. Meyer; p. 44, PI. 10 (fig. 1)

1857 - Eryon spinimanus Germar in Quenstedt; p. 805, PI. 99 (figs.

26-27)

1857 - Eryon speciosus Mùnster in Quenstedt; p. 806, PI. 90 (fig. 28,

31)

1862 -Eryon propinquus (Schlotheim) in Oppel; p. 12, PI. 1 (figs. 2-4),

PI. 2 (fig. 1)

1862 - Eryon spinimanus Germar in Oppel; p. 13, PI. 2 (fig. 2)

1 862 - Eryon Schuberti v. Meyer in Oppel; p. 1 8, PI. 3 (fig. 5)

1 885 - Eryon spinimanus Germar in Quenstedt; p. 408, PI. 32 (figs.

4-5)

1 904 - Eryon propinquus (Schlotheim) in Peiser; p. 25

1904 - Eryon propinquus (Schlotheim) in Walther; p. 173

1907 - Eryon propinquus (Schlotheim) in v. Knebel; p. 213, PI. 13

1908 - Eryon propinquus (Schlotheim) in Engel; p. 470

1822 - E/yon propinquus (Schlotheim) in Van Straelen; p. 1224

1825 - Coleia propinqua (Schlotheim) in Van Straelen; p. 148, Text-

fig. 67

1904 - Eryon spinimanus Germar in Walther; p. 1 73

1 904 - Eryon Schuberti v. Meyer in Walther; p. 1 73

1 904 - Eryon Schuberti v. Meyer in Peiser; p. 34

1907 - Eryon spinimanus Germar in v. Knebel; p. 217, Text-fig. 1,

PI. 15

1907 - Eryon Schuberti v. Meyer in v. Knebel; p. 219

1 908 - Eryon spinimanus Germar in Engel; p. 470

1924 - Eryon propinquus (Schlotheim) in Balss; p. 174, Figs. 1, 7

1925 - Coleia spinimana (Germar) in Van Straelen; p. 150, PI. 5

1925 - Knebelia Schuberti (v. Meyer) in Van Straelen; p. 123

1928 - Eryon propinquus (Schlotheim) in Beurlen; p. 225

1929- Coleia propinqua (Schlotheim) in Glaessner; p. 127

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART 1

25

1929 - Coleia spinimanus (Germar) in Glaessner; p. 128

1929 - Knebelia Schuberti (v. Meyer) in Glaessner; p. 228

1930 - Cyclocaris propinqua (Schlotheim) in Beurlen & Glaessner;

Text-fig. 12

1934 -Eryon propinquus (Schlotheim) in Dacqué; PI. 42 (fig. 2)

1968 - Eryon propinquus (Schlotheim) in Leich; p. 90

1969 - Cycleryon propinquus (Schlotheim) in Glaessner; R 470, Text-

fig. 274/4a, b

1969 - Knebelia schuberti (v. Meyer) in Malz; p. 296, Text-fig. 6

1969 - Cycleryon propinquus (Schlotheim) in Forster; p. 55, PI. 3 (fig. 1 )

1969 - Cycleryon propinquus (Schlotheim) in Malz; p. 293, Text-fig. 3

1978 - Cycleryon propinquus (Schlotheim) in Barthel; p. 328, 330, PI.

44-45

1994 - Cycleryon propinquus (Schlotheim) in Frickhinger; p. 116, Figs.

197-198

1 994 - Cycleryon spinimanus (Germar) in Frickhinger; Text-fig. 1 99

1994 - Knebelia schuberti (v. Meyer) in Frickhinger; p. 120, Fig. 2 1 0

1 995 - Cycleryon propinquus (Schlotheim) in Dietl, Kapitzke & Rieter;

PI. 2 (fig. 2)

2000 - Cycleryon propinquus (Schlotheim) in Ròper, Rothgaenger &

Rothgaenger; Text-fig. 16

2001 - Cycleryon spinimanus (Germar) in Schweigert; p. 5, Text-fig. 1,

PI. 1 (figs. 1-2), PI. 2 (figs. 1-2), PI. 3 (figs. 1-4)

2001 - Cycleryon spinimanus (Germar) in Dietl & Schweigert; p. 108,

Text-fig. 140

Diagnosis: carapace subcircular ventrally flattened;

median postcervical carina strong and branchial carinae

weak; cervical and postcervical incisions deep dividing

margin into three parts; pereiopods I-IV chelate; pereio-

pod V achelate; pereiopod I larger and stronger than

pereiopods II-V; pereiopods successively shorter posteri-

orly; uropodal exopod petaloid without diaeresis.

Lectotype: MNHB MB. A. 1107 (Schlotheim, 1822

- PI. 3, fig. 2), housed in thè Museum fur Naturkunde der

Humboldt-Universitàt in Berlin.

Our studies revealed that Eryon spinimanus from

Solnhofen and figured by Walch (1773 - PI. 14, fig. 1) is

synonym with Cycleryon propinquus, as discussed below.

The originai specimen is lost, so we decide to designate

thè specimen BSPG AS I 982 as thè neotype (= allotype

of Cycleryon propinquus) of Germar ’s species, housed in

thè Bayerische Staatssammlung fur Palàontologie und

Geologie in Munich.

Our studies revealed that thè minute Eryon schuberti

ffom Solnhofen and figured by v. Meyer (1836 - PI. 12,

figs. 3, 6) is synonym with Cycleryon propinquus, as dis¬

cussed below. This specimen, SMF X/m 144-146, is thè

holotype of v. Meyer’s species, housed in Natur-Museum

Senckenberg in Frankfurt am Main.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - Tithonian).

Type locality: Solnhofen.

Material: Oppel ( 1 862) reported 36 specimens belong-

ing to this species, so divided: five from thè Redenbacher

collection, two from thè Schlotheim collection, four from

thè Fraas collection and 25 from thè collection of thè Bay¬

erische Staatssammlung fur Palàontologie und Geologie

in Munich. We studied of this originai sample two speci¬

mens of Schlotheim collection, housed in thè Museum fur

Naturkunde der Humboldt-Universitàt in Berlin, and three

specimens, housed in thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich in which we iden-

tified one specimen, illustrated by Munster (1839), and

two specimens, illustrated by Oppel (1862). We report thè

list of thè specimens, studied by Munster and Oppel:

BSPG AS VII 765 - Munster (1839), PI. 3 (fig. 1)

- length of thè specimen: 10 cm

BSPG AS V 31 - Oppel (1862), PI. 1 (fig. 2)

- length of thè specimen: 12 cm

BSPG AS VI 42 - Oppel (1862), PI. 2 (fig. 1)

- length of thè specimen: 10 cm

Oppel (1862) reported four specimens which he

assigned to Cycleryon spinimanus (Germar, 1827) so

divided: one from thè Alberti collection (today SMNS),

one from thè collection of thè Bayerische Staatssa¬

mmlung fur Palàontologie und Geologie in Munich

and two from thè Fraas collection (today SMNS). We

studied two specimens of this originai sample, housed

in thè Bayerische Staatssammlung fur Palàontologie und

Geologie in Munich in which we identified one specimen,

illustrated by Oppel (1862), and one specimen, illustrated

by v. Knebel (1907). We report thè list of thè specimens,

studied by Oppel and v. Knebel:

BSPG AS VI 40 - Oppel (1862), PI. 2 (fig. 2)

- length of thè specimen: 12 cm

BSPG AS I 982 - Knebel (1907), PI. 15 (fig. 1)

- length of thè specimen: 10 cm

Oppel (1862) reported 56 specimens which he

assigned to Eryon schuberti v. Meyer, 1836, so divided:

16 from thè Redenbacher collection and 40 from thè

collection of thè Bayerische Staatssammlung tur Palà¬

ontologie und Geologie in Munich. We studied of this

originai sample four specimens, housed in thè Bayer¬

ische Staatssammlung fìir Palàontologie und Geologie in

Munich illustrated by Munster (1839). We report thè list

of thè specimens, studied by Munster:

BSPG AS VII 751 - Munster (1839), PI. 7 (fig. 6)

- length of thè specimen: 2 cm

BSPG AS VII 755 - Munster (1839), PI. 7 (fig. 2)

- length of thè specimen: 2 cm

BSPG AS VII 756 - Munster (1839), PI. 7 (fig. 7)

- incomplete specimen

BSPG AS VII 757 - Munster (1839), PI. 7 (fig. 8)

- length of thè specimen: 1.5 cm

Description. Large eryonid with densely tuberculate

exoskeleton.

Carapace. Carapace subcircular ventrally flattened,

wider than long. Frontal margin strengthened with small

tuburcles, without rostrum. Cervical and postcervical

incisions deep dividing margin into three parts. Ocular

incision small and narrow. Median postcervical carina

strongly tuberculate. Branchial carinae weak and tuber¬

culate. Posterior margin of carapace with row of strong

tubercles. Lateral margins of carapace strongly dentate.

Dorsal surface of carapace densely tuberculate.

Abdomen. Somite I subrectangular smaller than thè

others. Somites II-VI subrectangular of equal length.

Pleurae and lateral parts of somites II-VI fìnely tubeculate,

median part of somites smooth. Pleurae strongly dentate.

Somites I-V with a strongly raised median tergal carina.

Telson lanceolate with a strongly raised basai tubercle,

a pair of proximal tubercles dose to lateral margins, a

median longitudinal carina, two submedian carinae, and

rounded distai extremity. Protopod subsquare. Uropodal

endopod subrectangular with a strong median longitudi¬

nal carina. Uropodal exopod petaloid without diaeresis

and with a strong median longitudinal carina.

26

ALESSANDRO GARASSINO & GONTER SCHWEIGERT

Cephalic appendages. Eyestalk very short. 3rd maxil-

liped elongate and spineless. Antennular articles thin of

which thè distai one carries two short multiarticulate

flagella of equal length. Antennal articles short and stout

of which thè distai one carries a thin multiarticulate flag-

ellum. Mouth fìeld displaying crista dentata on 3rd maxil-

liped and mandibles (Fig. 5).

Fig. 5 - Cycleryon propinquus (Schlotheim, 1822), ventral view of

carapace, line drawing (F. Fogliazza).

Thoracic appendages. Pereiopod I with strong and

elongate propodus. Chela of pereiopod I with fixed finger

gently curved in thè distai part and movable finger curved

distally with enlarged distai extremity. Thin median longi-

tudinal carina extends for total length of propodus. Inner

margins of movable and fixed fìngers edentate in thè male;

three strong proximal teeth in thè inner margin of movable

tinger in thè temale (Figs. 5, 6). Surface of merus, carpus,

propodus, movable and fixed fìngers smooth. Pereiopods

’l-'V with small chelae of equal length. Outer margins

of carpus and propodus of pereiopods II-III strengthened

w ith a row of small tubercles. Pereiopods V shorter than

others and achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. loday, Cycleryon Glaessner, 1965,

known from thè Middle Jurassic (Callovian) to thè Upper

Jurassic ( lithonian) of Europe (France and Germany),

includes five species. One from thè Middle Jurassic, C.

giganteus (Van Straelen, 1923) (France - Callovian);

three from thè Upper Jurassic (Kimmeridgian - Titho-

nian) of Germany, C. propinquus (Schlotheim, 1822), C.

elongatus (Miinster, 1 839), and C. orbiculatus (Miinster,

1839). Cycleryon armatus (v. Knebel, 1907) and C. sub-

rotundus are interpreted as synonyms with C. orbiculatus

(see C. armatus and C. subrotundus in this paper).

Recently, Garassino & Schweigert (2004) described

from thè Upper Jurassic (Tithonian) of Blumenberg

quarry, dose to thè city of Eichstàtt (Bavaria, S Ger¬

many), thè new species C. wulfi.

Fig. 6 - Cycleryon spinimanus (Germar, 1827), pereiopod I, line draw¬

ing (F. Fogliazza).

Germar (1827) pointed out that thè presence of two

strong spines in thè proximal part of thè inner margin of

movable finger was a morphological character enough

to distinguish C. spinimanus (Germar, 1827) from C.

propinquus (Schlotheim, 1822). Subsequently, after thè

review of Cycleryon Glaessner, 1965, in thè Nusplingen

Lithographic Limestone (S Germany) by Schweigert

(200 la), C. spinimanus is considered as thè female of C.

propinquus.

The poor state of preservation of thè type speci¬

men of Eryon schuberti v. Meyer, 1836, did not allow

extending its morphological analysis. In fact, it is very

ditficult to recognize some diagnostic characters in order

to describe this species, as already reported by Oppel

(1862) who pointed out thè poor preservation of thè

specimen. V. Meyer (1836) erected E. schuberti as new

species of Eryon Desmarest, 1822. Van Straelen (1925)

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART 1

27

ascribed this species to Knebelia Van Straelen, 1922,

for thè small size of its specimen with strongly curved

chelae of perieopod I. Even though thè originai speci¬

men shows thè frontal margin with two subrectangular

lobes typical of Knebelia , this morphological characters

is also observable in many eryonid juvenile specimens

belonging to C. propinquus. We consider thè resemblace

between K. schuberti and K. bilobata , type species of

thè genus, only a homoemorphy in order to consider thè

originai specimen of K. schuberti as juvenile stage of C.

propinquus.

Cycleryon orbiculatus (Munster, 1839)

Figs. 7a, 10, PI. XVI (figs. 1, 2, 3, 4)

1839 - Eryon orbiculatus Munster; p. 7, PI. 5 (figs. 1-2)

1839 - Eryon latus Munster; p. 8, PI. 5 (figs. 4-6), PI. 7 (fig. 1)

1839 - Eryon subrotundus Munster; p. 12, PI. 7 (figs. 4-5), nov. syn.

1853 - Eryon orbiculatus Munster in Frischmann; p. 27

1853 - Eryon latus Munster in Frischmann; p. 27

1853 - Eryon subrotundus Miinster in Frischmann; p. 27

1862 - Eryon orbiculatus Munster in Oppel; p. 14, PI. 2 (fig. 3)

1 862 - Eryon subrotundus Munster in Oppel; p. 14

1904 - Eryon orbiculatus Munster in Walther; p. 173

1907 - Eryon orbiculatus Munster in v. Knebel; p. 215

1907 - Eryon armatus v. Knebel; p. 215, PI. 14, nov. syn.

1925 - Coleia orbiculata (Munster) in Van Straelen; p. 147

1925 - Coleia armata (Knebel) in Van Straelen; p. 153

1928 - Eryon orbiculatus Munster in Beurlen; p. 277

1929 - Coleia orbiculata (Munster) in Glaessner; p. 127

1929 - Coleia armata (Knebel) in Glaessner; p. 125

1994 - Cycleryon orbiculatus (Munster) in Frickhinger; p. 116, Fig.

196

2001 - Cycleryon orbiculatus (Munster) in Dietl & Schweigert; p. 54,

Text.fig. 89

Diagnosis: carapace subcircular ventrally flattened;

median postcervical carina, and branchial carinae absent;

cervical and postcervical incisions deep dividing margin

into three parts; suborbitai tooth strong and outer orbitai

tooth strong; pereiopods I-IV chelate; pereiopod V ache-

late; pereiopod I larger and stronger than pereiopods II-V;

pereiopods successively shorter posteriorly; uropodal

exopod petaloid without diaeresis.

Lectotype: BSPG AS VII 760 - Munster (1839), PI.

5 (fig. 2), housed in thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich.

Our studies revealed that Eryon armatus from an

unknown locality and fìgured by v. Knebel (1907 - PI.

14) is synonym with Cyclery’on orbiculatus , as discussed

below. This specimen, MNHB MB. A. 236, is thè lecto¬

type of v. KnebeTs species, housed in thè Museum fur

Naturkunde der Humboldt-Universitàt in Berlin.

Our studies revealed that Eryon subrotundus from

Eichstàtt and figured by Munster (1839 - PI. 7, Fig. 5)

is synonym with Cycleryon orbiculatus , as discussed

below. This juvenile specimen, BSPG AS VII 762, is

thè lectotype of Miinster’s species, housed in thè Bayer¬

ische Staatssammlung fur Palàontologie und Geologie in

Munich.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - lower Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported 40 specimens

belonging to this species, so divided: 15 from thè Reden-

bacher collection, 24 from thè collection of thè Bayer¬

ische Staatssammlung fUr Palàontologie und Geologie

in Munich and 1 from thè Palaeontological Museum of

Zurich (from Nusplingen). We studied three specimens of

this originai sample, housed in thè Bayerische Staatssa¬

mmlung fur Palàontologie und Geologie in Munich

in which we identified three specimens, illustrated by

Munster (1839), and one specimen, illustrated by Oppel

(1862). We report thè list of thè specimens, studied by

Munster and Oppel:

BSPG AS VI 38 - Munster (1839), PI. 7 (fig. 1)

- length of thè specimen: 8 cm

BSPG AS VII 760 - Munster (1839), PI. 5 (fig. 2)

- length of thè specimen: 6 cm

BSPG AS I 990 - Oppel (1862), PI. 2 (fig. 3)

- length of thè specimen: 5 cm

V. Knebel (1907) did not report in thè description of

E. armatus thè number of studied specimens, illustrat-

ing only one specimen, today housed in thè Museum fur

Naturkunde der Humboldt-Universitàt in Berlin:

MNHB MB. A. 236 - v. Knebel (1907), PI. 14

- length of thè specimen: 4 cm

Munster (1839) reported two specimens which he

assigned to Eryon subrotundus Munster, 1839, belonging

to thè collection of thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich where, today, they

are stili preserved. We report thè list of thè specimens,

studied by Munster:

BSPG AS VII 762 - Miinster (1839), PI. 7 (figs. 5)

- length of thè specimen: 3.5 cm

BSPG AS VII 763 - Munster (1839), PI. 7 (fig. 4)

- incomplete specimen

Description. Median-sized eryonid with densely

tuberculate exoskeleton.

Carapace. Carapace subcircular ventrally flattened,

wider than long. Frontal margin without rostrum. Cervical

and postcervical incisions deep dividing margin into three

parts. Ocular incision small and narrow. Suborbitai tooth

strong. Median postcervical and branchial carinae absent.

Lateral margins of carapace strongly dentate. Dorsal sur-

face of carapace densely tuberculate.

Abdomen. Somite I subrectangular smaller than thè

others. Somites II- VI subrectangular of equal length

and with fìnely tubeculate dorsal surface. Pleurae finely

tuberculate. Somites I-V with a weakly raised median

tergal carina. Telson laceolate with a median longitudi-

nal carina, two submedian carinae, and rounded distai

extremity. Protopod subsquare. Uropodal endopod subre¬

ctangular with a strong median longitudinal carina. Uro¬

podal exopod petaloid without diaeresis and with a strong

median longitudinal carina.

Cephalic appendages. Eyestalk very short. 3rd maxil-

liped elongate and spineless. Antennular articles thin of

which thè distai one carries two short multiarticulate flag¬

ella of equal length. Antennal articles short and stout of

which thè distai one carries a thin multiarticulate flagellum.

Thoracic appendages. Pereiopod I with strong and

stout propodus. Chela of pereiopod I with movable finger

curved at distai extremity and straight fixed fìnger. Thin

median longitudinal carina extends for total length of

propodus. Inner margins of movable and fixed fingers

edentate. Surface of merus, carpus, propodus, movable,

28

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

and fixed fingers smooth. Pereiopods II-IV with small

chelae of equal length. Pereiopods V shorter than others

and achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. Miinster (1839) described this species on

eight specimens, pointing out that they differed from thè

type species because thè chela of pereiopod I was shorter

and thicker. Today, only three originai specimens are

available and their study allowed recognition of other

morphological characters useful to justify thè systematic

validity of this species. In fact, thè strong suborbitai tooth,

thè strong outer orbitai tooth and thè lack of carinae on thè

dorsal surface of carapace distinguish C. orbiculatus from

thè type species and from C. elongatus (see C. elongatus

in this paper) (Fig. 7).

Fig. 7 - a) Cycleryon orbiculatus (Miinster, 1839), line drawing; b)

Cycleryon elongatus (Miinster, 1839), line drawing (F. Fogliazza).

V. Knebel (1907) described E. armatus on some small-

sized specimens, pointing out that they differed from C.

propinquus and C. orbiculatus for thè shorter chela of

pereiopod I, thè triangular shape of ocular incision, and

thè shorter abdomen. The study of thè only available

specimen allowed recognition of some morphological

characters, typical of C. orbiculatus in order to consider

thè studied specimen as juvenile stage of this species. In

fact, thè shape of carapace wider than long, thè strong

suborbitai tooth, thè strong outer orbitai tooth, and thè

lack of carinae on thè dorsal surface of carapace are char¬

acters observable also in Miinster’s species.

Miinster (1839) reported only some morphological

characters of carapace and pereiopod I of Eryon sub-

rotundus because thè poor state of preservation of thè

specimens. Oppel (1862) pointed out that thè originai

specimens studied by Miinster could be juvenile stages

of C. orbiculatus. This is confirmed by thè re-study of thè

lectotype using ultraviolet illumination.

Cycleryon elongatus (Miinster, 1839)

Figs. 7b, 10, PI. XVI (fig. 5)

1839 - Eryon elongatus Miinster; p. 9, PI. 5 (fig. 7)

1 853 - Eryon elongatus Miinster in Frischmann; p. 27

1 862 - Eryon elongatus Miinster in Oppel; p. 15, PI. 2 (fig. 4)

1904 -Eryon elongatus Miinster in Walther; p. 173

1907 - Eryon elongatus Mùnster in v. Knebel; p. 216

1 925 - Coleia elongata (Miinster) in Van Straelen; p. 1 53

1929 - Coleia elongata (Miinster) in Glaessner; p. 126

1969 - Cycleryon elongatus (Miinster) in Malz; p. 295, Text-fig. 4

1999 - Cycleryon elongatus (Miinster) in Frickhinger; p. 116, Fig. 195

Diagnosis: carapace subcircular ventrally flattened;

median postcervical carina and branchial carinae weak;

cervical and postcervical incisions deep dividing margin

into three parts; pereiopods I-IV chelate; pereiopod V

achelate; pereiopod I larger and stronger than pereiopods

II-V; pereiopods successively shorter posteriorly; uropo-

dal exopod rounded without diaeresis.

Lectotype: BSPG AS VII 761 (Miinster, 1839 — PI.

5, fig. 7), housed in thè Bayerische Staatssammlung fur

Palàontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported three specimens

belonging to this species and housed in thè collection of

thè Bayerische Staatssammlung fur Palàontologie und

Geologie in Munich. We studied this originai sample

recognizing one specimen, illustrated by Miinster (1839),

and two specimens, illustrated by Oppel (1 862). We report

thè list of thè specimens, studied by Miinster and Oppel:

BSPG AS VII 761 - Miinster (1839), PI. 5 (fig. 7)

- length of thè specimen: 8 cm

BSPG AS VI 43 - Oppel (1862), PI. 2 (fig. 4)

- length of thè specimen: 8 cm

BSPG AS I 939 - Oppel (1862), PI. 2 (fig. 4)

- length of thè specimen: 8 cm

One specimen (n. cat. 9419 - Wulf collection) was

used to describe thè carapace, poorly preserved in thè

originai specimens.

Description. Median-sized eryonid with densely

tuberculate exoskeleton.

Carapace. Carapace subcircular ventrally flattened,

wider than long. Frontal margin without rostrum. Cervical

and postcervical incisions deep dividing margin into three

parts. Ocular incison small and narrow. Median postcer¬

vical carina weakly tuberculate and raised. Branchial

carinae weak and tuberculate. Lateral margins of cara¬

pace weakly dentate. Dorsal surface of carapace densely

tuberculate.

Abdomen. Somite I subrectangular smaller than thè

others. Somites II- VI subrectangular of equal length.

Pleurae and lateral parts of somites II-VI finely tubercu¬

late, median part of somites smooth. Pleurae weakly den¬

tate. Somites I-V without a median tergal carina. Telson

lanceolate with two submedian carinae and rounded distai

extremity. Protopod subsquare. Uropodal endopod subre¬

ctangular with a strong median longitudinal carina. Uro¬

podal exopod rounded without diaeresis and with a strong

median longitudinal carina.

Cephalic appendages. Eyestalk very short. 3rd maxil-

liped elongate and spineless. Antennular articles thin of

which thè distai one carries two short multiarticulate

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

29

flagella of equal length. Antennal articles short and stout

of which thè distai one carries a thin multiarticulate flag¬

ellimi. Mouth field displaying crista dentata on 3rd maxil-

liped and mandibles.

Thoracic appendages. Pereiopod I with strong, stout

propodus. Chela of pereiopod I with movable finger

curved at distai extremity and straight fixed finger. Thin

median longitudinal carina runs for total length of pro¬

podus. Inner margins of movable and fixed fingers eden-

tate. Surface of merus, carpus, propodus, movable, and

fixed fingers smooth. Pereiopods II-IV with small chelae

of equal length. Pereiopods V shorter than others and

achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. Even though Miinster (1839) did not

establish thè morphological characters, typical of this

species, he pointed out that thè short chela of pereiopod

I was similar between C. elongatus and C. orbiculatus.

Oppel (1862) compared C. elongatus with C. propinquus,

telling that thè sizes of pereiopod I and of thè abdomen

are different between thè two species. In fact, pereiopod I

is longer and thè abdomen is smaller in thè type species.

Even though there are some morphological affinities

among C. elongatus, C. orbiculatus, and C. propinquus,

thè study of thè originai specimens revealed that thè tail

fan in C. elongatus is completely different than that on thè

other two. In fact, thè telson in C. elongatus is character-

ized by only two lateral carinae, while in thè other two

species, thè telson has one median longitudinal carina

and two lateral carinae. Moreover, thè uropodal exopod

in C. elongatus has a rounded shape with rounded distai

extremity, while in thè other two species, thè uropodal

exopod has a petaloid shape with pointed distai extremity

at thè conjunction between thè lower and lateral margins

(Fig.7). ‘

Genus Eryon Desmarest, 1 822

Type species: Macrourites arctiformis Schlotheim,

1822

Eryon arctiformis (Schlotheim, 1820)

Fig. 11, PI. VII, PI. XVII (fig. 1)

1820 - Macrourites arctiformis Schlotheim; p. 37

1822 - Macrourites arctiformis Schlotheim; p. 34, PI. 3 (fig. 1)

1822 - Eryon Cuvieri Desmarest in Brogniart; p. 128, PI. 10 (fig. 4)

1823 - Eryon Cuvieri Desmarest in Kriiger; p. 587

1825 - Eryon Cuvieri Desmarest; p. 209, PI. 34 (fig. 3)

1825 - Eryon Cuvieri Desmarest in Kriiger; p. 265

1827 - Eryon Cuvieri Desmarest in Germar; p. 98

1827 - Eryon acutusl Germar; p. 100

1836 -Eryon Cuvieri Desmarest in v. Meyer; p. 273, PI. 12 (fig. 5)

1837 - Eryon arctiformis (Schlotheim) in Bronn; p. 474, PI. 137 (fig.

2)

1839 -Eryon arctiformis (Schlotheim) in Miinster; p. 3, PI. 1

1839 - Eryon pentagonus Miinster; p. 10, PI. 6 (fig. 1)

1839 -Eryon subpentagonus Miinster; p. 10, PI. 6 (fig. 2)

1862 - Eryon arctiformis (Schlotheim) in Oppel; p. 15, PI. 3 (fig. 1)

1904 - Eryon arctiformis (Schlotheim) in Peiser; p. 25

1907 - Eryon arctiformis (Schlotheim) in v. Knebel: p. 200, PI. 1 1 (fig.

1,6), PI. 12

1924 - Eryon arctiformis (Schlotheim) in Balss; p. 174, Text-figs. 3, 6

1925 - Eryon arctiformis (Schlotheim) in Van Straelen; p. 114, Text-

fig. 63

1929 -Eryon arctiformis (Schlotheim) in Glaessner; p. 164

1969 - Eryon arctiformis (Schlotheim) in Glaessner; R 470, Text-fig.

274/2

1969 - Eryon arctiformis (Schlotheim) in Malz; p. 292, Text-figs. 1-2

1994 - Eryon arctiformis (Schlotheim) in Frickhinger; p. 120, Figs.

203-204

2005b - Eryon arctiformis (Schlotheim) in Schweigert & Garassino;

p. 498

Diagnosis: carapace subcircular ventrally flattened;

median postcervical carina and branchial carinae weak;

cervical and postcervical incisions deep dividing margin

into three parts; pereiopods I-IV chelate; pereiopod V

achelate; pereiopod I larger and stronger than pereiopods

II-V; pereiopods successively shorter posteriorly; uropo¬

dal exopod petaloid without diaeresis.

Holotype: Macrourites arctiformis (= Eryon arcti¬

formis), MNHB K 38 MB.A. 253 (Schlotheim, 1822 - PI.

3, fig. 1; length of thè specimen: 7.5 cm), housed in thè

Museum fur Naturkunde der Humboldt-Universitàt in

Berlin.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported 170 specimens

belonging to this species, so divided: 80 from thè Reden-

bacher collection and 90 from thè collection of thè Bayer-

ische Staatssammlung fur Palaontologie und Geologie in

Munich. We studied of this originai sample thè holotype

by Schlotheim, housed in thè Museum fur Naturkunde

der Humboldt-Universitàt in Berlin, and six specimens,

housed in thè Bayerische Staatssammlung fur Palàontolo-

gie und Geologie in Munich in which we identified five

specimens, illustrated by Miinster (1839), and one speci¬

men, illustrated by Oppel (1862). We report thè list of thè

specimens, studied by Miinster and Oppel:

BSPG AS VII 758 - Miinster (1839), PI. 2

- length of thè specimen: 16 cm

BSPG AS VII 764 - Miinster (1839), PI. 1 (fig. 2)

- length of thè specimen: 1 1 cm

BSPG AS VII 766 - Miinster (1839), PI. 1 (fig. 3)

- length of thè specimen: 10 cm

BSPG AS VII 767 - Miinster (1839), PI. 1 (fig. 1)

- length of thè specimen: 12 cm

BSPG AS VII 1431 - Miinster (1839), PI. 6 (fig. 2)

- length of thè specimen: 7 cm

BSPG AS VI 41 - Oppel (1862), PI. 3 (fig. 1)

- length of thè specimen: 10 cm

Description. Large eryonid with densely tuberculate

exoskeleton.

Carapace. Carapace subcircular ventrally flattened,

wider than long. Frontal margin strengthened with small

tuburcles and without rostrum. Cervical and postcervical

incisions deep dividing margin into three parts. Ocular

incisions developed in conical structures carrying an elon-

gate eyestalk. Median postcervical carina strongly tuber¬

culate and raised. Branchial carinae weak and tuberculate.

Posterior margin of carapace concave. Lateral margins of

carapace strongly dentate, originally hearing fringes simi¬

lar to those of M. longimanatus (not figured in thè recon-

struction). These fringed margins are only observable in

30

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

rapidly buried specimens, and they are not preserved in

moults. Dorsal surface of carapace densely tuberculate.

Abdomen. Somite I subrectangular smaller than thè

others. Somites II-VI subrectangular of equal length and

with finely tuberculate dorsal surface. Pleurae strongly

dentate. Somites II-VI with a strongly raised median

tergal carina. Telson lanceolate with a strongly raised

median longitudinal carina and with pointed distai

extremity. Protopod subsquare. Uropodal endopod sub¬

rectangular with a strong median longitudinal carina.

Uropodal exopod petaloid without diaeresis and with a

strong carina extending parallel to outer margin. In rap¬

idly buried specimens, thè uropodal endopod and exopod

show a fringed margin.

Cephalic appendages. Eyestalk elongate. 3rd maxil-

liped elongate and spineless. Antennular articles thin

of which thè distai one carries two short multiarticulate

flagella of equal length. Antennal articles short and stout

of which thè distai one carries a thin multiarticulate flag-

ellum. Mouth fìeld displaying crista dentata on 3rd maxil-

liped and mandibles.

Thoracic appendages. Pereiopod I with strong and

elongate propodus. Chela of pereiopod I with fìxed finger

straight and movable fìnger bent distally with enlarged

distai extremity. Inner margins of movable and fìxed

fingers edentate. Surface of merus, carpus, propodus,

movable, and fìxed fingers smooth. Pereiopods II-IV with

small chelae of equal length. Pereiopods V shorter than

others and achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. Today, Eryon Desmarest, 1822, known

from thè Middle Jurassic (Callovian) to thè Upper Cre-

taceous (Campanian) of Europe (Germany, France and

England), includes five species. Three from thè Middle

Jurassic: E. perroni Etallon, 1859, and E. ellipticus Van

Straelen, 1922 (France - Callovian), and E. sublevis

Carter, 1886 (England - Oxfordian). One from thè Upper

Jurassic: E. arctiformis (Schlotheim, 1822) (Germany

- Kimmeridgian/Tithonian). One from thè Lower Creta-

ceous: 1E. neocomiensis Woodward, 1881 (Silesia -Neo-

comian; inclusion in Eryon is very uncertain). Eryon

yehoachi Remy & Avnimelech, 1955, from thè Upper

Cretaceous of Israel was redescribed as a fossil stomato-

pod species by Hof (1998).

Genus Rosenfeldia Garassino, Teruzzi &

Dalla Vecchia, 1996

Type species: Rosenfeldia triasica Garassino, Teruzzi

& Dalla Vecchia, 1996

Rosenfeldia oppeli (Woodward, 1866)

PI. XVII (fig. 2)

1866 - Eryon Oppeli Woodward; p. 500, PI. 24 (fig. 4)

1925 - Eryon Oppeli Woodward in v. Straelen; p. 120

1929 - Eryon Oppeli Woodward in Glaessner; p. 166

1963 - Eryon oppeli Woodward in Kuhn; p. 22

2003 - Rosenfeldia oppeli (Woodward) in Schweigert & Garassino; p.

178, Fig. 2A

2004a - Rosenfeldia oppeli (Woodward) in Schweigert, p. 70, Text-fig.

without number

2004b - Rosenfeldia oppeli (Woodward) in Schweigert; p. 329, Text-

fig. without number

2005a - Rosenfeldia oppeli (Woodward) in Schweigert & Frattigiani;

p. 1 98, Text-fig. without number

2005b - Rosenfeldia oppeli (Woodward) in Schweigert & Frattigiani; p.

328, Text-fig. without number

Holotype by monotypy: BM 44886, housed in thè

Naturai History Museum in London.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: originally labelled as coming from Sol- .

nhofen. According to thè lithology of thè rock matrix it

comes from thè surroundings of Eichstàtt.

Material: Woodward (1866) reported one incomplete

specimen, belonging to thè Hàberlein collection and

today housed in thè Naturai History Museum in London.

In 2004 a second specimen was recovered from thè Mòm-

sheim Formation of thè Horstberg quarry near Mòmsheim

(H. & K. Schumacher collection).

Discussion. Woodward (1866) based his description

on an incomplete specimen belonging to thè Hàberlein

collection, justifìng thè institution of Eryon oppeli on

two morphological characters: pereiopod I is small if

compared with thè four succeding pereiopods and smaller

than pereiopod I present in thè other species of Eryon

Desmarest, 1822, and thè subrectangular telson exhibited

rounded lateral margins that are different from thè sub-

triangular telson typical of Eryon. Even though thè poor

state of preservation of thè specimen did not allow rec-

ognition of thè main morphological characters useful to

establish its placement in this genus, surely thè specimen

must be ascribed to thè family Eryonidae De Haan, 1841,

for thè lack of thè diaeresis on thè uropodal exopod.

Garassino et al. (1996) described Rosenfeldia from

thè Late Triassic (Norian) of NE Italy, belonging to thè

family Eryonidae for thè lack of a diaeresis on thè uropo¬

dal exopod and characterized by a subrounded carapace,

chela of pereiopod I with movable finger longer than fìxed

finger and curved at thè distai extremity, subrectangular

telson with dentate lateral margins, and uropodal endopod

and exopod with dentate lateral margins.

Even though thè originai specimen by Woodward

is incomplete (only maxillae, pereiopods I-V, abdomi¬

nal somites IV-VI and tail fan are preserved), its study

pointed out some morphological characters, such as thè

dentate pleural margins of somites IV-VI, thè dentate lat¬

eral margins of thè telson and thè dentate lateral margins

of uropodal endopod and exopod, typical of Rosenfeldia.

Therefore, E. oppeli is referred to this genus. In one of thè

newly recovered specimens (H. & K. Schumacher collec¬

tion) thè carapace is also missing, and its overall preser-

vational state is worse than that of thè holotype. Another

two specimens (Frattigiani collection) exhibit also poor

remains of thè carapace (Schweigert, 2005; Schweigert

& Garassino, in press). The ascription of E. oppeli to this

genus expands thè stratigraphic range of Rosenfeldia.

Rosenfeldia is now represented by two species, R.

triasica (Norian) and R. oppeli (Tithonian). Rosenfeldia j

was only known by thè type species R. triasica Garassino,

Teruzzi & Dalla Vecchia, 1996, from thè Upper Triassic

(Norian) of NE Italy. R. oppeli differs from thè Triassic

one in hearing smaller chelae in pereiopod I, thè lateral

margins of pereiopods I-IV being smooth and thè telson

having a weak longitudinal median carina.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

31

Genus Knebelia Van Straelen, 1 922

Type species: Eryon bilobatus Miinster, 1839

Knebelia bilobata (Miinster, 1839)

Fig. 11, PI. Vili, PI. XVII (figs. 3, 4)

1839 - Eryon bilobatus Miinster; p. 11, PI. 6 (figs. 3-5)

1 853 - Eryon bilobatus Miinster in Frischmann; p. 27

1862 - Eryon bilobatus Miinster in Oppel; p. 16, PI. 3 (fig. 2)

1904 - Eryon bilobatus Miinster in Peiser; p. 25, 32

1904 - Eryon bilobatus Miinster in Walther; p. 173

1907 - Miinsteria bilobata (Miinster) in v. Knebel; p. 223

1922 - Knebelia bilobata (Miinster) in Van Straelen; p. 982

1925 - Knebelia bilobata (Miinster) in Van Straelen; p. 121

1929 - Knebelia bilobata (Miinster) in Glaessner; p. 227

1969 - Knebelia bilobata (Miinster) in Glaessner; R 470, Text-fig.

274/1

1994 -Knebelia bilobata (Miinster) in Frickhinger; p. 120, Fig. 209

Diagnosis: carapace subsquare ventrally flattened;

frontal margin with two subrectangular lobes and with a

deep and very narrow cephalic incision; cervical and post-

cervical incisions weak dividing margin into three parts;

cervical groove weak; pereiopods I-IV chelate; pereiopod

V achelate; pereiopod I larger and stronger than pereio¬

pods II-V; pereiopods successively shorter posteriorly;

uropodal exopod without diaeresis.

Lectotype: BSPG AS VI 39 (Oppel, 1862 - PI. 3, fig.

2), housed in thè Bayerische Staatssammlung fur Palàon-

tologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported ten specimens

belonging to this species, so divided: four from thè

Redenbacher collection and six from thè collection of thè

Bayerische Staatssammlung fur Palàontologie und Geolo¬

gie in Munich. We studied four specimens of this originai

sample, housed in thè Bayerische Staatssammlung tur

Palaontologie und Geologie in Munich in which we iden-

tified three specimens illustrated by Miinster (1839), and

one specimen, illustrated by Oppel (1862). We report thè

list of thè specimens, studied by Miinster and Oppel:

BSPG AS VII 752 - Miinster (1839), PI. 6 (fig. 5)

- length of thè specimen: 3 cm

BSPG AS VII 753 - Miinster (1839), PI. 6 (fig. 4)

- length of thè specimen: 7 cm

BSPG AS VII 754 - Miinster (1839), PI. 6 (fig. 3)

- length of thè specimen: 7 cm

BSPG AS VI 39 - Oppel (1862), PI. 3 (fig. 2)

- length of thè specimen: 8 cm

Description. Large eryonid with densely tuberculate

exoskeleton.

Carapace. Carapace subsquare ventrally flattened.

Frontal margin formed with two subrectangular lobes

with a deep and very narrow cephalic incision and with¬

out rostrum. Cervical and postcerical incisions weak

dividing margin into three parts. Ocular incision narrow

located laterally. Cervical groove weak. Posterior margin

narrow and strongly convex and strengthened by a row of

small tubercles. Dorsal surface densely tuberculate.

Abdomen. Somite I subrectangular smaller than thè

others. Somites II-VI subrectangular of equal length and

with finely tuberculate dorsal surface. Pleurae smooth.

Somites II-VI with a median tergal carina strongly raised.

Telson laceolate with one longitudinal median carina, two

slightly raised submedian carinae, one basai tubercle,

and pointed distai extremity. Protopod subsquare. Uro¬

podal endopod subrectangular with a strong longitudinal

median carina. Uropodal exopod subrectangular without

diaeresis and with a strong longitudinal median carina.

Cephalic appendages. Eyestalk short. 3rd maxilliped,

antennulae and antennae poorly preserved.

Thoracic appendages. Pereiopod I with slender and

elongate propodus. Lower margin of merus with a row

of small spines. Chela of pereiopod I with fixed finger

straight and movable finger longer than fixed finger and

bent distally with pointed distai extremity. Inner margins

of movable and fixed fingers edentate. Surface of merus,

carpus, propodus, movable, and fixed fingers smooth.

Pereiopods II-IV with small chelae of equal length. Perei¬

opods V shorter than others and achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. Today, Knebelia Van Straelen, 1922,

known only in thè Upper Jurassic (Tithonian) of Ger-

many, includes thè only species, K. bilobata (Miinster,

1839). Eryon schuberti v. Meyer, 1836, often included

in Knebelia , represents a juvenile specimen of Cycleryon

propinquus (Schlotheim, 1822).

Family Polychelidae Wood-Mason, 1874

Genus Palaeopentacheles v. Knebel, 1907

Type species: Eryon roettenbacheri Miinster, 1839

Palaeopentacheles roettenbacheri (Miinster, 1839)

Fig. 11, PI. IX, PI. XVII (fig. 5), PI. XVIII (figs. 1, 2, 3, 4)

1839 -Eryon Rottenbacheri Miinster; p. 13, PI. 7 (fig. 10)

1 848 - Eryon Rottenbacheri Miinster in Bronn; p. 468

1852 - Eryon Rottenbacheri Miinster in Quenstedt; p. 267

1853 - Eryon Redenbacheri Miinster in Frischmann; p. 27

1856 - Eryon Redenbacheri Miinster in v. Meyer; p. 49, PI. 9 (figs.

1-3)

1862 - Eryon Redenbacheri Miinster in Oppel; p. 18, PI. 3 (figs. 6-7)

1904 - Eryon Redenbacheri Miinster in Peiser; p. 43

1904 -Eryon Redenbacheri Munster in Walther; p. 173

1907 - Palaeopentacheles Redenbacheri (Munster) in v. Knebel; p.

226, Text-fig. 2

1924 - Eryon Redenbacheri Miinster in Balss; p. 181, Text-fig. 9

1925 - Palaeopentacheles Redenbacheri (Munster) in Van Straelen;

p. 124

1929 - Palaeopentacheles Redenbacheri (Munster) in Glaessner; p. 299

1969 - Palaeopentacheles roettenbacheri (Munster) in Glaessner; R

471, Text-fig. 275/2

1969 - Palaeopentacheles redenbacheri (Munster) in Malz; p. 298,

Text-fig. 5

1 969 - Palaeopentacheles redenbacheri (Munster) in Forster; p. 47

1994 - Palaeopentacheles redenbacheri (Munster) in Frickhinger; p.

122, Fig. 220

1994- Palaeopolycheles longipes (Fraas) in Frickhinger; p. 122, Fig. 22 1

2001 - Palaeopolycheles roettenbacheri (Miinster) in Dietl & Sch-

weigert; p. 71

2005b - Palaeopolycheles roettenbacheri (Munster) in Schweigert &

Garassino; p. 498

32

ALESSANDRO GARASSINO & GÙNTER SCHWEIGERT

Diagnosis: carapace subrectangular ventrally flat-

tened; projecting front having two strong and elongate

teeth with serrate outer margins; lateral margins convex

and denticulate; cervical groove well marked; median

postcervical carina weak; branchial carinae strong;

median postrostral carina weak; cervical and postcervical

incisions absent; pereiopods I-IV chelate; pereiopod V

achelate; pereiopod I larger and stronger than pereiopods

II-V; pereiopods successively shorter posteriorly; uropo-

dal exopod without diaeresis.

Lectotype: BSPG AS I 993 (Oppel, 1862 - PI. 3, fìg.

6), housed in thè Bayerische Staatssammlung fur Palàon-

tologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported 20 specimens

belonging to this species, so divided: six from thè Reden-

bacher collection and 14 from thè collection of thè Bay¬

erische Staatssammlung fur Palàontologie und Geologie

in Munich. We studied four specimens of this originai

sample, housed in thè Bayerische Staatssammlung fìir

Palàontologie und Geologie in Munich in which we iden-

tified two specimens, illustrated by v. Meyer (1856) and

two specimens, illustrated by Oppel (1862). We report thè

list of thè specimens, studied by Meyer and Oppel:

BSPG AS I 991 - v. Meyer (1856), PI. 9 (fìg. 2)

- incomplete specimen

BSPG AS I 992 - v. Meyer (1856), PI. 9 (fìg. 1)

- length of thè specimen: 5 cm

BSPG AS I 989 - Oppel (1862), PI. 3 (fìg. 7)

- incomplete specimen

BSPG AS I 993 - Oppel (1862), PI. 3 (fìg. 6)

- length of thè specimen: 4 cm

Description. Small-sized eryonid with densely tuber-

culate exoskeleton.

Carapace. Carapace subrectangular ventrally flattened.

Projecting front having two strong and elongate teeth with

serrate outer margins. Cervical and postcervical incisions

absent. Ocular incision deep located anteriorly. Cervical

groove well marked. Median postcervical carina weak

and tuberculate. Branchial carinae strong and tuberculate.

Median postrostral carina weak. Lateral margins convex

and denticulate. Posterior margin strongly convex. Dorsal

surface densely tuberculate.

Abdomen. Somite I subrectangular smaller than thè

others. Somites II-VI subrectangular of equal length and

with finely tuberculate dorsal surface. Pleurae pointed

and tuberculate. Somites II-VI with a strongly raised

median tergal carina, hearing a strong spine on each

pleura directed backward. Somites II-V with a strong

spine directed outward partially to cover each pleura.

Telson laceolate with two submedian tuberculate carinae

strongly raised, one strong basai spine, and tridentate

distai extremity. Protopod subsquare. Uropodal endopod

subrectangular with a strong longitudinal median carina.

Uropodal exopod subrectangular with one small spine in

thè middle of thè lateral margin and without diaeresis.

Cephalic appendages. Antennular articles thin of

which thè distai one carries two short multiarticulate

flagella of equal length. Antennal articles short and stout

ot which thè distai one carries a thin multiarticulate flag-

ellum. Scaphocerite spatolate with a row of small spine

along outer margin.

Thoracic appendages. Pereiopod I with slender and

elongate merus with a row of small spines along thè lower

margin and strong propodus with upper and lower margins

hearing small tubercles. Chela of pereiopod I with slender

and pointed fixed finger and movable finger curved dis-

tally. Inner margins of movable and fixed fingers with

strong teeth increasing in size distally. Surface of merus,

carpus, propodus, movable, and fixed fingers tuberculate.

Pereiopods II-IV with small chelae of equal length. Perei¬

opods V shorter than thè others and achelate.

Abdominal appendages. Pleopods not preserved.

Discussion. V. Knebel (1907) established thè new

genus Palaeopentacheles with thè type species P. roet-

tenbacheri (Miinster, 1839) from thè Upper Jurassic

(Tithonian) of Germany.

Recently, thè discovery of P. starri by Schweitzer

& Feldmann (2001) from thè Oligocene of Washington

(United States) largely expands thè geographic and strati¬

graphic range of this genus.

Family Palinuridae Latreille, 1802

Genus Palinurina Miinster, 1839

Type species: Palinurina longipes Miinster, 1839

Included species: Palinurina longipes Miinster, 1839;

Palinurina tenera Oppel, 1862

Palinurina longipes Miinster, 1839

Fig. 12, PI. XVIII (fig. 5), PI. IXX (fig. 1)

1839 - Palinurina longipes Miinster; p. 37, PI. 14 (fig. 8)

1839 - Palinurina intermedia Miinster; p. 37, PI. 14 (figs. 9,10?), PI.

29 (Fig. 8), nov. syn.

1 839 - Palinurina pygmea Miinster; p. 38, PI. 14 (fig. 11), nov. syn.

1862 - Palinurina longipes Miinster in Oppel; p. 86, PI. 24 (fig. 1)

1 862 - Palinurina pygmea Miinster in Oppel; p. 87, PI. 24 (fig. 2)

1 873 - Palinurina longipes Miinster in Seebach; p. 345

1873 - Palinurina tenera Oppel in Seebach; p. 345

1889 - Palinurina pygmea Miinster in Oppenheim; p. 718

1904 - Palinurina longipes Miinster in Walther; p. 174

1904 - Palinurina pygmea Miinster in Walther; p. 174

1925 - Palinurina longipes Miinster in Van Straelen; p. 223

1929 - Palinurina longipes Miinster in Glaessner; p. 301

1969 - Palinurina longipes Miinster in Glaessner; R 474, Text-fig.

277/6

1973 - Palinurina longipes Miinster in Forster; p. 38, Text-Figs. 4-7

1994 - Palinurina longipes Miinster in Frickhinger; p. 125, Fig. 222

2001 - Palinurina longipes Miinster in Dietl & Schweigert; p. 71

2005b - Palinurina longipes Miinster in Schweigert & Garsssino; p. 498

Diagnosis: carapace subrectangular ventrally flat¬

tened; rostrum short, shorter than supraorbital spine;

cervical and postcervical grooves deep; hepatic and

branchiocardiac grooves thin and weak; supraorbital and

antennal spines well developed; suborbitai spine small;

abdomen longer than carapace; pereiopod I shorter and

stronger than pereiopods II-V; well-developed articles of

thè antennae; very elongate flagellum of antennae; uropo¬

dal exopod without diaeresis.

Lectotype: BSPG AS V 46 (Oppel, 1862 - PI. 24, fig.

1), housed in thè Bayerische Staatssammlung fìir Palàon¬

tologie und Geologie in Munich.

I

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

33

Our studies revealed that Palinurìna intermedia from

Solnhofen and figured by Miinster (1839 - PI. 14, figs.

9-10) is synonym with Palinurina longipes , as discussed

below. The specimen, BSPG AS VII 768, is thè lectotype

of Miinster’s species, housed in thè Bayerische Staatssa-

mmlung fìir Palàontologie und Geologie in Munich.

Our studies revealed that Palinurina pygmea from

Solnhofen and figured by Miinster (1839 - PI. 14, fig. 11)

is synonym with Palinurina longipes , as discussed below.

This specimen, BSPG AS I 996, is thè lectotype of Miin-

ster’s species, housed in thè Bayerische Staatssammlung

tur Palàontologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (upper Kim-

meridgian - Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported 28 specimens

belonging to this species, so divided: six from thè Reden-

bacher collection and 22 from thè collection of thè Bay¬

erische Staatssammlung fìir Palàontologie und Geologie

in Munich. We studied two specimens of this originai

sample, housed in thè Bayerische Staatssammlung fìir

Palàontologie und Geologie in Munich in which we iden-

tified one specimen, illustrated by Miinster (1839) and

one specimen, illustrated by Oppel (1862). We report thè

list of thè specimens, studied by Miinster and Oppel:

BSPG AS V 47 - Miinster (1839), PI. 14 (fig. 8)

- length of thè specimen: 3 cm

BSPG AS V 46 - Oppel (1862), PI. 24 (fig. 1)

- length of thè specimen: 4 cm

Miinster (1839) reported a sample of three specimens

which he assigned to Palinurina intermedia. We studied of

this originai sample two specimens, housed in thè Bayer¬

ische Staatssammlung fìir Palàontologie und Geologie in

Munich illustrated by Miinster (1839). We report thè cata-

logue number of thè two specimens, studied by Miinster:

BSPG AS VII 767 - Miinster (1839), PI. 14 (fig. 10)

- length of thè specimen: 2 cm

BSPG AS VII 768 - Miinster (1839), PI. 14 (fig. 9)

- length of thè specimen: 3 cm

Oppel (1862) reported a sample of 48 specimens

which he assigned to Palinurina pygmea Miinster, 1839,

so divided: 12 from thè Redenbacher collection and 36

from thè collection of thè Bayerische Staatssammlung

tur Palàontologie und Geologie in Munich. We studied of

this originai sample one specimen, housed in thè Bayer¬

ische Staatssammlung tur Palàontologie und Geologie in

Munich illustrated by Oppel (1862). We report thè cata-

logue number of thè specimen, studied by Oppel:

BSPG AS I 996 - Oppel (1862), PI. 24 (fig. 2)

- length of thè specimen: 2 cm

One specimen (n. cat. 0315 - Wulf collection) was

used to describe thè carapace which was poorly preserved

in thè originai sample.

Description. Small-sized palinurid with finely tuber-

culate exoskeleton.

Carapace. Carapace subrectangular ventrally flat-

tened, something missing. Rostrum short, shorter than thè

supraorbital spines. A ridge of two/three spines directed

forward thè supraorbital spine. Antennal spine well devel-

oped, longer than thè supraorbital spine. A ridge of three

spines directed forward thè antennal spine. Suborbitai

spine small, forming a ridge with other two small spines

directed forward thè ocular incision. Postcervical and cer-

vical grooves deep. Hepatic and branchiocardiac grooves

thin and weak. Two rows of small tubercles extend par¬

afici on thè median dorsal line of carapace, between thè

postcervical and branchiocardiac grooves.

Abdomen. Somites I-VI of equal length. Somites II-

V with denticulated margins and with a sharp terminal

spine. Tergal and pleural surfaces with small tubercles.

Telson subrectangular with rounded distai extremity and

with a short spine located in thè median part of lateral

margin. Uropodal endopod, exopod, and telson of equal

length. Uropodal endopod and exopod with rigid and flex-

ible portions. Uropodal exopod without diaeresis.

Cephalic appendages. Antennular articles thin of

which thè proximal one is longer than thè other two.

Antennal articles strong and stout with denticulate lateral

margins. Antennal flagellum can exceed five time thè

length of thè body.

Thoracic appendages. Pereiopod I shorter and stronger

than pereiopods II-V. Segments of pereiopods I-V with

small tubercles arranged in parallel rows.

Abdominal appendages. Pleopods with an unseg-

mented peduncle which carries two elongate multiarticu-

late flagella.

Discussion. Palinurina Miinster, 1839, belongs to thè

family Palinuridae Latreille, 1802, known from thè Upper

Triassic with Archaeopalinurus Pinna, 1974, from thè

Norian of Cene (Bergamo, N Italy) (Pinna, 1974). Three

genera belong to this family from thè Jurassic: Astacodes

Bell, 1863 (Germany, Slovakia - Kimmeridgian), Pa-

laeopalinurus Bachmayer, 1954 (Germany - Tithonian),

and Palinurina Miinster, 1839 (England - Sinemurian/

Toarcian; Germany - Tithonian). Five genera belong to

this family from thè Cretaceous: Astacodes Bell, 1863

(Europe, United States, Australia - Turonian/Santonian),

Eurycarpus Schliiter, 1 868 (Germany - Senonian), Jasus

Parker, 1 883 (Lebanon - Cenomanian), Linuparus White,

1847 (Germany, England, France, Canada, Antarctic

Peninsula, United States, Mexico and New Zealand

- Turonian/Campanian), and Palinurus Weber, 1795

(Italy -Albian; Germany, Lebanon - Turonian/Senonian).

Finally, three genera belong to this family from thè Ceno-

zoic: Archaeocarabus McCoy, 1849 (England - Lute-

tian), Jasus Parker, 1883 (New Zealand - Oligocene), and

Justitia Holthuis, 1946 (Italy - Lutetian).

Among thè above mentioned genera, Palinurina Miin-

ster, 1 839, strongly resembles, for thè generai shape of thè

body, Archaeopalinurus. However, if we examine more

deeply thè two forms, it is possible to point out some mor-

phological differences that justify thè separation into two

different genera. If we compare our specimens with thè

reconstruction available in thè litterature, particularly that

concemig A levis (Pinna, 1974 - PI. 15, fig. 3; Garassino

& Teruzzi, 1993 - Fig. 33) we can notice remarkable

differences in thè shape and in thè omamentation of thè

tail fan, in thè length of pereiopods (thè II and thè IV are

longer than thè others in A. levis, while II-V have equal

length in P. longipes), in thè length of thè antennal flagel¬

lum which is five time longer than thè body in thè German

species. Other diagnostic differences appear to be seen

in thè morphology and omamentation of thè antennal

peduncles and of thè carapace, but thè latter is too poorly

preserved in A. levis to be sure about that.

Palinurina longipes Miinster, 1839, was thè subject

of a review by Forster (1973). Today, Palinurina, known

from thè Sinemurian to thè Tithonian (Lower-Upper

34

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

Jurassic) of Europe (England and Germany), includes two

species, P. longipes and P. tenera (see next description).

Miinter (1839) described Paìinurina intermedia

on some morphological characters different to those

observed in thè other species of thè same genus, in partic-

ular P. longipes : thè very elongate antennae are smooth,

lacking thè fine lateral setae, thè large thick antennal

stalk has three spiny articulations and all pereiopods are

relatively shorter and thicker than those of thè type spe¬

cies.

However Oppel (1862) reported that thè specimens

studied by Munster and labelled as P. intermedia were

extremely poorly preserved and thè generai outline of thè

body and thè pereiopods did not provide any differences

with that of P longipes, so considering Munster ’s species

as synonym with thè type species.

Munster ( 1 839) described Paìinurina pygmea on some

morphological characters different to those observed in

thè other species of thè same genus, in particular P. lon¬

gipes : thinner and shorter antennae, shorter and more

curved pereiopods and shorter abdomen. However, thè

study of thè only survived specimen, belonging to this

species, has pointed out some morphological characters,

similar to those observed in P longipes (cfr. description of

P. longipes), such as shorter and stronger pereiopod I, seg-

ments of pereiopods I-V with small tubercles arranged in

two parallel rows and antennal articles with small tuber¬

cles and spiny lateral margins. Therefore, w e consider P.

pygmea as synonym of P. longipes, type species of this

genus on thè basis of thè above mentioned morphological

characters, common between thè two species. Probably,

thè specimens belonging to P pygmea, gathered by Mun¬

ster in thè same species on thè basis of thè same size, were

juvenile stages of P. longipes.

Paìinurina tenera Oppel, 1 862

PI. IXX (figs. 2, 3)

1862 - Paìinurina tenera Oppel; p. 86, PI. 23 (fig. 4)

1 873 - Paìinurina tenera Oppel in Seebach; p. 345

1889 - Paìinurina tenera Oppel in Oppenheim; p. 718

1904 - Paìinurina tenera Oppel in Walther; p. 1 74

2005b - Paìinurina tenera Oppel in Schweigert & Garassino; p. 498

The poor state of preservation of thè originai incom¬

plete specimen by Oppel has led to thè research of new

specimens from private collections in order to emend

thè morphological description of Paìinurina tenera. One

specimen (n. cat. SMNS 65476 - Frattigiani collection)

preserved in ventral view allowed recognition of some

morphological characters in order to deepen thè descrip¬

tion of this species. However, thè preservation in ventral

view of Frattigiani ’s specimen did not pennit description

of thè carapace and tail fan.

Diagnosis: abdomen longer than carapace; pereiopod

1 shorter and stronger than pereiopods II-V; articles of thè

antennae well developed; flagellum of antennae short;

uropodal exopod without diaeresis.

Lectotype: BSPG AS I 995 (Oppel, 1862 - PI. 23, fig.

4), housed in thè Bayerische Staatssammlung fur Palàon-

tologie und Geologie in Munich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Eichstàtt.

Material: Oppel (1862) reported 25 specimens

belonging to this species and belonging to thè collection

of thè Bayerische Staatssammlung fìir Palàontologie und

Geologie in Munich. We know of this originai sample one

specimen and illustrated by Oppel (1862). We report thè

catalogue number of thè specimen, studied by Oppel:

BSPG AS I 995 - Oppel (1862), PI. 23 (fig. 4)

- incomplete specimen

Description. Small-sized palinurid with fìnely tuber-

culate exoskeleton.

Carapace. Not preserved.

Abdomen. Somites I-VI of equal length. Somites II-

V with denticulated margins and with a sharp terminal

spine. Tergal and pleural surfaces with small tubercles.

Uropodal exopod without diaeresis.

Cephalic appendages. Antennular articles thin of which

thè proximal one is longer than thè other two. Antennal

articles strong and stout with smooth lateral margins.

Antennal flagellum as long as thè length of thè body.

Thoracic appendages. Pereiopod I shorter and stronger

than pereiopods II- V. Segments of pereiopods I-V with

rare small tubercles not arranged in parallel rows.

Abdominal appendages. Not preserved.

Discussion. Oppel (1862) justified thè difference of

P. tenera from thè type species based upon two morpho¬

logical characters: thè antennal flagellum which is shorter

than that of P longipes and thè pereiopods that are longer

than those of thè type species. Even though it is difficult

to compare thè two species for their rarity, we are able

to confimi thè systematic validity of Oppel’s species not

only by thè above-mentioned characters, but also for

thè antennal articles which have smooth lateral margins

(denticulate in thè type species) and for thè segments of

pereiopods I-V with rare small tubercles not arranged in

parallel rows (many tubercles arranged instead in parallel

rows in thè type species).

Family Cancrinidae Beurlen, 1930

Genus Cancrinos Munster, 1839

Type species: Cancrinos claviger Munster, 1839

Cancrinos claviger Munster, 1839

Fig. 12, PI. X, PI. IXX (fig. 4), PI. XX (figs. 1, 2, 3, 4, 5)

1839 - Cancrinos claviger Munster; p. 43, PI. 15 (fig. 1)

1839 - Cancrinos latipes Munster; p. 44, PI. 15 (fig. 2), nov. syn.

1862 - Cancrinus claviger Munster in Oppel; p. 88, PI. 24 (fig. 3)

1862 - Cancrinus latipes Munster in Oppel; p. 88, PI. 24 (fig. 4)

1904 - Cancrinos claviger Munster in Walther; p. 174

1904 - Cancrinos latipes Munster in Walther; p. 174

1925 - Cancrinos claviger Munster in Van Straelen; p. 226, Text-fig.

108

1929 - Cancrinos claviger Munster in Glaessner; p. 109

1929 - Cancrinos latipes Munster in Glaessner; p. 109

1969 - Cancrinos claviger Munster in Glaessner; R 475, Text-fig.

280/2

1994 - Cancrinos claviger Munster in Frickhinger; p. 115, Figs. 193-

194

1999 - Cancrinos claviger Munster in Frickhinger; p. 42, Fig. 66

1999 - Cancrinos latipes Munster in Frickhinger; p. 42, Fig. 67

2006 - Cancrinos claviger Munster in Garassino & Schweigert; p. 75

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

35

This species is very rare in thè lithographic lime-

stone and usually thè specimens are not well preserved.

However, we were able to give its diagnosis and morpho-

logical description by thè identification of some morpho-

logical characters of carapace, abdomen, pereiopods, and

cephalic appendages in thè originai specimens of this spe¬

cies. Moreover, one specimen (BSPG 1975 IX 13) housed

in thè Bayerische Staatssammlung fur Palàontologie und

Geologie in Munich was used to describe for thè fìrst time

thè shape of thè abdominal somites and tail fan, not pre¬

served in thè originai sample.

Diagnosis: carapace subrectangular ventrally flat-

tened; rostrum absent; cervical groove deep; abdomen as

long as carapace; pereiopod I shorter and stronger than

pereiopods II-V; petaloid flagellimi of thè antennae strong

and stout; uropodal exopod without diaeresis.

Neotype: BSPG AS I 998 (Oppel, 1862 - PI. 24, figs.

5, 6), housed in thè Bayerische Staatssammlung fiir Pala-

ontologie und Geologie in Munich. According to Munster

(1839) thè holotype was purchased by Prof. v. Breda from

Leiden. Today thè specimen seems to be lost, because

it was not traceable either in thè Teyler’s Museum in

Harlem, or in thè British Naturai History Museum in

London, where specimens of v. Breda collection is said

to be housed.

Our studies revealed that Cancrinos latipes from

Moritzbrunn near Eichstàtt and figured by Munster (1839

- PI. 15, fig. 2) is synonym with Cancrinos claviger, as

discussed below. This specimen, BSPG AS VII 769, is thè

holotype of Miinster’s species, housed in thè Bayerische

StaatssammlungfurPalàontologieundGeologieinMunich.

Stratigraphic range: Upper Jurassic (Tithonian).

Type locality: Solnhofen.

Material: Oppel (1862) reported three specimens

belonging to this species and belonging to thè collec¬

tion of thè Bayerische Staatssammlung tur Palàontologie

und Geologie in Munich where, today, they are stili pre¬

served.

BSPG AS I 997 - Oppel (1862), PI. 24 (fìg. 3)

- length of thè specimen: 10 cm

BSPG AS I 998 - Oppel (1862), PI. 24 (figs. 5, 6)

- incomplete specimen

BSPG AS I 999 - Oppel (1862), PI. 24 (fig. 4)

- length of thè specimen: 10 cm

Munster (1839) reported one specimen which he

assigned to Cancrinos latipes Munster, 1839, belonging

to thè collection of thè Bayerische Staatssammlung fìir

Palàontologie und Geologie in Munich where, today, it is

stili preserved.

BSPG AS VII 769 - Munster (1839), PI. 15 (fig. 2)

- incomplete specimen

Description. Large palinurid with strongly tubercu-

late exoskeleton.

Carapace. Carapace subrectangular ventrally flat-

tened. Rostrum absent. Cervical groove deep, located in

thè median part. Posterior margin strengthened by a thin

marginai carina.

Abdomen. Somites I-VI subrectangular of equal

length. Pointed pleurae with a strong, sharp tooth and

denticulate margin. Telson subrectangular with rounded

distai extremity. Uropodal endopod and exopod with

rigid and flexible portions. Rigid portion of exopod finely

tuberculate. Uropodal exopod without diaeresis.

Cephalic appendages. Antennular flagella thin and

elongate. Antennal articles strong and stout with denticu¬

late lateral margins. Antennal flagellum short and petaloid

with 22 segments narrowing distally.

Thoracic appendages. Pereiopod I shorter and stronger

than pereiopods II-V with a strong and stout dactylus.

Upper and lower margins of I-V strengthened by a row of

strong spines directed forward.

Abdominal appendages. Not preserved.

Discussion. Today, Cancrinos Munster, 1839, known

only from thè Upper Jurassic (Tithonian) of Germany

includes just one species, C. claviger , very rare in thè

lithographic limestone. Recently, thè new species C.

libanensis has been recorded from thè Cenomanian of

Lebanon, extending thè stratigraphic range of this genus

(Garassino & Schweigert, 2006).

Oppel (1862), revising thè two species distinguished

by Munster (1839), C. claviger and C. latipes, used as

diagnostic character thè different number of segments of

thè antennal flagellum. In fact, thè author pointed out that

thè antennal flagellum had 15 or 16 segments in C. clavi¬

ger, while it had 18 segments in C. latipes. In reality, thè

study of thè originai specimens of thè two species pointed

out that thè antennal flagellum exhibits thè same number

of segments (ca. 22 in all), as also confirmed by thè study

of new specimens, housed in thè Bayerische Staatssa¬

mmlung tur Palàontologie und Geologie in Munich, in

which thè number of segments is included between 21

and 23, thè exact number not easily countable because of

thè telescoping in thè distai part of thè flagellum. More¬

over, thè pereiopod I is shorter than thè others in both

species. Finally, we consider C. claviger as type species

of Cancrinos because thè most important diagnostic char¬

acters that identify thè genus are preserved in thè originai

specimens of this species.

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

Eryma modestiforme

Palaeastacus fuciformis

7,

Fig. 8 - Summarizing figure ot thè astacideans preserved in Solnhofen fauna, line drawing (F. Fogliazza).

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

37

Etallonia longimana

Fig. 9 - Summarizing figure of thè astacideans and thalassinideans preserved in Solnhofen fauna, line drawing (F. Fogliazza).

ALESSANDRO GARASSINO & GONTER SCHWEIGERT

Cycleryon propinquus

Cycleryon orbiculatus

Cycleryon elongatus

Fig. 10 - Summarizing figure of palinurids preserved in Solnhofen fauna, line drawing (F. Fogliazza).

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART 1

Eryon arctiformis

Fig. 1 1 - Summarizing figure of palinurids preserved in Solnhofen fauna, line drawing (F. Fogliazza).

40

ALESSANDRO GARASSINO & GONTER SCHWEIGERT

Palinurina longipes

Cancrinos claviger

Fig. 12 - Summarizing figure of palinurids preserved in Solnhofen fauna, line drawing (F. Fogliazza).

ACKNOWLEDGEMENTS

We wish to thank M. Nose, Bayerische Staatssa-

mmlung fur Palaontologie und Geologie, Munich, for

permitting study of thè historic decapod collection, A.

Ross, Department of Palaeontology, Naturai History

Museum, London, for permitting study of thè holotype

of Eryon oppeli, C. Neumann, Museum tur Naturkunde

der Humboldt-Universitàt, Berlin, for thè pictures of thè

holotypes of thè Schlotheim collection, T. Jellinek, For-

schungsinstitut and Naturmuseum Senckenberg (Frank¬

furt am Main) for thè picture of thè holotype of Eryon

schuberti, N. Hauschke, Halle/Saale for searching for thè

holotypes of Mecochirus bajeri and Eryma leptodacty-

lina, M. Wulf, H. Tischlinger, P. Riidel and R. Frattigiani

for permitting study of some specimens of their private

collections, R. M. Feldmann, Geology Department, Kent

State University (Ohio) and G. Teruzzi, Museo Civico di

Storia Naturale di Milano, for thè useful advice in thè

draft of thè manuscript, careful review and criticism.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

41

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Museo civico di Storia naturale di Milano, Milano.

Schweigert G., Garassino A., Hall R. L., Hauff R. B. & Karasawa H.,

2003 - The lobster genus Uncina Quenstedt, 1851 (Crustacea:

Decapoda: Astacidea: Unicinidae) from thè Lower Jurassic.

Stuttgart er Beitràge zur Naturkunde, ser. B, Stuttgart, 332: 1-43.

Schweigert G., Garassino A. & Riou B., 2006 - First record of

Stenochirus Oppel, 1861 (Crustacea: Decapoda: Stenochiridae)

from thè Callovian (Middle Jurassic) of La Voulte sur-Rhóne.

Neues Jahrbuch fur Geologie und Palàontologie, Monatshefte,

Stuttgart, 2006 (2): 65-77.

Schweigert G., Krishna J., Pandey B. & Pathak D. B., 1996 - A new

approach to thè correlation of thè Late Kimmeridgian Beckeri

Zone across thè Tethyan Sea. Neues Jahrbuch fur Geologie und

Palàontologie, Abhandlungen, Stuttgart, 202: 345-373.

Schweigert G. & Ròper M., 2001 - Neue Krebse der Gattung Palaeast-

acus (Crustacea: Decapoda: Erymidae) aus oberjurassischen Plat-

tenkalken Suddeutschlands. Stuttgarter Beitràge zur Naturkunde,

ser. B, Stuttgart, 313: 1-10.

Schweitzer C. & Feldmann R. M., 2001 - New Cretaceous and Tertiary

decapod crustaceans from western North America. Bulletin of thè

Mizunami Fossil Museum, Mizunami, 28: 173-210.

Secretan S., 1964 - Les crustacés décapodes du Jurassique supérieur et

du Crétacé de Madagascar. Mémoires Muséum national d’Histoire

naturelle, Paris, ser. C, 15: 1-224.

Seebach K. von, 1873 - Ueber fossile Phyllosomen von Solnhofen.

Zeitschrift der Deutschen Geologischen Gesellschaft, Berlin, 25:

340-346.

Simpson M. I. & Middleton R., 1 985 -Gross morphology and thè mode of

life of two species of lobster from thè Lower Cretaceous of England:

Meyeria ornata (Phillips) and Meyerella magna (M’Coy). Transac¬

tion of thè Royal Society of Edinburgh, Edinburgh, 76: 203-215.

Stenzel H. B., 1945 - Decapod crustaceans from thè Cretaceous of

Texas. University of Texas, Austin, 4401 : 401-476.

Taylor B. J., 1979 - Macrurous Decapoda from thè Lower Cretaceous of

South-Eastem Alexander Island. British Antarctic Survey, 8 1 : 1-39.

Tshudy D. M. & Babcock L. E., 1 997 - Morphology-based phylogenetic

analysis of thè clawed lobsters (family Nephropsidae and thè new

family Chilenophoberidae). Journal of Crustacean Biology, 17:

257-267.

Van Straelen V., 1922 - Les Crustacés décapodes du Callovien de

La Voulte-sur-Rhòne (Ardèche). Comptes Rendus Académie des

Sciences, Paris, 175: 982-984.

Van Straelen V. 1923 - Description de Crustacés décapodes macroures

nouveaux des Terrains secondaires. Annales de la Société royale

Zoologique et Malacologique de Belgique, Bruxelles, 53 (1922),

1923: 84-93.

Van Straelen V., 1924-1925 - Contribution a l’étude des crustacés

décapodes de la Période Jurassique. Académie Royale de Belgique,

Cl. Se., Mém., Coll. 4, ser. 2, Bruxelles, 7 (1): 1-462.

Van Straelen V., 1927 - Contribution à Tétude des Crustacées

décapodes fossiles de la Péninsule Ibérique. Eos, Revista Espanola

d Entomologia, Madrid, 3: 79-94.

Van Straelen V., 1936 - Crustacés Décapodes nouveaux ou peu

connus de l’époque crétacique. Bulletin du Musée royal d’Histoire

naturelle de Belgique, Bruxelles, 12 (45): 1-50.

Walch J. E. I., 1773 - Naturgeschichte der Versteinerungen zur

Erlàuterung der Knorrischen Sammlung von Merkwiirdigkeiten

der Natur. Felfiecker, Numberg, 1: 1-187.

Walther J., 1 904 - Die Fauna der Solnhofener Plattenkalke, bionomisch

betrachtet . Jenaische Denkschriften, Jena, 11: 133-214.

Weber F., 1795 - Nomenclator entomologicus secundum entomologiam

systematicum ili. Fabricii, adjectis speciebus recens detectis et

varietatibus.

Winckler T. C., 1883 - Etudes carcinologique sur les genres Pemphix,

Glyphea et Araeosternus. Archives Musée Teyler, sér. 2, 1: 73-124.

White A., 1847 - Short descriptions of some new species of Crustacea

in thè collection of thè British Museum. Annals and Magazine of

Naturai History, London, 20 (132): 205-207.

Wood-Mason J., 1874 - On new or little-known Crustaceans. Proceed-

ings of thè Asiatic Society ofBengal, Calcutta, 63 (2): 230-232.

Woodward H., 1866 - Notes on thè species of thè genus Eryon

Desmarest, from thè Lias and Oolite of England and Bavaria.

Quarterly Journal of thè Geological Society, London, 22: 494-

502.

Woodward H., 1876 - On some New Macrurous Crustacea from thè

Kimmeridge Clay of thè Sub-Wealden Boring, Sussex, and from

Boulogne-sur-mer. Quarterly Journal of thè Geological Society,

London, 32 (1), 125: 47-50.

Woodward H., 1881 - Contribution to thè study of Fossil Crustacea.

Geological Magazine, London, 8: 529-534.

Woodward H., 1900 - Further Notes on Podophthalmous Crustaceans

from thè Upper Cretaceous Formation of British Columbia, etc.

Geological Magazine, London, new series, decade 4, 7: 433-435.

Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italia.

e-mail: a.garassino@tin.it

Gunter Schweigert - Staatliches Museum fur Naturkunde, Rosenstein 1, 7091 1 Stuttgart, Germany.

e-mail: schweigert.smns@naturkundemuseum-bw.de

1 he l pper Jurassic Solr.hoten decapod crustacean fauna: review of thè types from old descriptions. Part I.

Infraorders Astacidea, Thalassinidea, and Palinura

Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano

Volume XXXIV — Fascicolo I

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

45

PI. I - 1) Eryma modestiforme (Schlotheim, 1822); 2) Eryma veltheimii (MUnster, 1839); 3) Palaeastacus fuciformis (Schlotheim,

1822). Reconstructions F. Fogliazza.

46

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

pl- 11 1 * l>ustulina minuta (Schlotheim, 1822); 2) Pseudastacus pustulosus (Miinster, 1839). Reconstructions F. Fogliazza.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

47

PI. Ili - 1) Stenochirus angustus (Miinster, 1839); 2) Glyphea pseudoscyllarus (Schlotheim, 1822). Reconstructions F. Fogliazza.

48

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

1 1. 1\ 1 ) Glyphea tenuis Oppel, 1860; 2) Squamosoglyphea squamosa (Munster, 1839). Reconstructions F. Fogliazza.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

49

PI. V- 1 ) Mecochirus longimanatus (Schlotheim, 1822); 2) Magila latimana Miinster, 1839. Reconstructions F. Fogliazza.

50

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

PI. VI - Cyeleryon propinquus (Schlotheim, 1822). Reconstruction F. Fogliazza

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

51

PI. VII -Eryon arctiformis (Schlotheim, 1822). Reconstruction F. Fogliazza.

52

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

PI. Vili - Knebelia bilobata (Munster, 1839). Reconstruction F. Fogliazza

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

53

PI. IX - Palaeopentacheles roettenbacheri (Munster, 1839). Reconstruction F. Fogliazza.

54

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

PI. X - Cancrinos claviger Munster, 1839. Reconstruction F. Fogliazza.

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART 1

55

PI. XI - 1) Eryma modestiforme (Schlotheim, 1822), holotype, n. cat. K 37 MB (MNHB) (x 2.5); 2) Eryma leptodactylina (Germar,

1827), holotype by monotypy, without catalogue number (IGH) (x 2); 3, 4) Eryma veltheimii (Munster, 1839), holotype by monotypy,

n. cat. AS VII 186 (BSPG), part and counter-part (x 0.8); 5) Palaeastacus fuciformis (Schlotheim, 1822), holotype, n. cat. K 36 MB. A

251 (MNHB) (x 1.5); 6) Pustulina minuta (Schlotheim, 1822), holotype, n. cat. K 41 MB (MNHB) (x 3).

56

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

1 4

3 6

i i. X II - 1 ) / ustulina minuta (Schlotheim, 1 822), n. cat. AS Vili 78 (BSPG) (x 1 .2); 2) “ Glyphea ” verrucosa Munster, 1 839, lectotype,

n.cat. AS VII 182 (BSPG) (x2.5); 3) Pseudastacus pustulosus (Munster, 1839), lectotype, n. cat. AS I 670 (BSPG) (x 0.6); 4) Pseu-

dashiL us muensteri Oppel, 1862, holotype by monotypy, n. cat. AS I 672 (BSPG) (naturai size); 5) Stenochirus angustus (Munster,

; 839), holotype by monotypy, n. cat. AS VII 322 (BSPG) (naturai size); 6) Stenochirus angustus (Munster, 1839), n. cat. 9836 (Wulf

collection) (x 0.5).

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

57

PI. XIII - 1) Stenoehirus mayeri Oppel, 1862, lectotype, n. cat. AS I 673 (BSPG) (naturai size); 2) Stenochirus mayeri Oppel, 1862, n.

cat. AS I 673 (BSPG), detail of thè chela of pereiopod I (x 4); 3) Glyphea pseudoscyllarus (Schlotheim, 1822), holotype, n. cat. K 43

MB. A. 255 (MNHB) (naturai size); 4) Glyphea tenuis Oppel, 1860, holotype by monotypy, n. cat. AS VIII 95 (BSPG) (naturai size);

5) Squamosoglyphea squamosa (Miinster, 1839), holotype by monotypy, n. cat. AS VII 734 (BSPG) (naturai size); 6) Mecochirus

longimanatus (Schlotheim, 1822), neotype, n. cat. AS V 45 (BSPG) (x 0.3).

58

ALESSANDRO GARASSINO & GÙNTER SCHWEIGERT

; ™ ' 1 ) Mecochirus brevimanus (Munster, 1839), lectotype, n. cat. AS VII 739 (BSPG) (x 0.7); 2) Mecochirus dubius (Munster,

- \ ec,otype; n‘ cat- AS 1 979 (BSPG) (x 1.5); 3) Etallonia longimana (Munster, 1839), lectotype, n. cat. AS VII 259 (BSPG) (x

u n8 te;™™ Master, 1839, lectotype, n. cat. AS VI 26 (BSPG) (x 2.5); 5) Magila latimana Munster, 1839, n. cat. 0117

1 1 tollection) (naturai size); 6) Magila desmarestii (Munster, 1839), holotype by monotypy, n. cat. AS VII 199 (BSPG) (naturai

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

59

PI. XV - 1) Magila robusta Oppel, 1862, holotype by monotypy, n. cat. AS I 675 (BSPG) (x 2); 2) Orhomalus deformis (Oppel, 1862),

holotype by monotypy, n. cat. AS Vili 115 (BSPG) (naturai size); 3) Cycleryon propinquus (Schlotheim, 1822), lectotype, n. cat.

MB. A. 1107 (MNHB) (naturai size); 4) Cycleryon spinimanus (Germar, 1827), neotype, n. cat. AS I 982 (BSPG) (x 0.6); 5) Cycleryon

spinimanus (Germar, 1827), neotype, n. cat. AS I 982 (BSPG), detail of thè pereiopod I (naturai size); 6) Eryon schuberti v. Meyer,

1836, holotype, n. cat. SMF X/m 144-146 (SMF) (x 2).

60

ALESSANDRO GARASSINO & GÙNTER SCHWEIGERT

I. \\ i - 1 1 Cyc leryon orbiculatus (Munster, 1839), lectotype, n. cat. AS VII 760 (BSPG) (x 0.8); 2) Cycleryon subrotundus (Miinster,

1 83^), lectotype, n. cat. AS VII 762 (BSPG) (x 1 .2); 3) Cycleryon subrotundus (Munster, 1 839), lectotype, n. cat. AS VII 762 (BSPG)

in ultraviolet ìllumination (x 1.2); 4) Cycleryon armatus (v. Knebel, 1907), lectotype, n. cat. MB. A. 236 (MNHB) (x 0 7)- 5 Cycleryon

elongatus (Munster, 1839), lectotype, n. cat. AS VII 761 (BSPG) (x 1.2).

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

61

PI. XVII - 1 Eryon arctiformis (Schlotheim, 1822), holotype, n. cat. K 38 MB. A. 253 (MNHB) (x 1.3); 2) Rosenfeldia oppeli (Wood-

ward, 1866), holotype by monotypy, n. cat. 44886 (BM) (x 2); 3) Knebelia bilobata (Miinster, 1839), lectotype, n. cat. AS VI 39

(BSPG) (x 0.8); 4) Knebelia bilobata (Miinster, 1839), n. cat. AS VII 754 (BSPG), detail of thè frontal margin (x 2.5); 5) Palaeopen-

tacheles roettenbacheri (Miinster, 1839), lectotype, n. cat. AS I 993 (BSPG) (x 1.2).

62

ALESSANDRO GARASSfNO & GUNTER SCHWEIGERT

PI. XVIII - 1) Palaeopentacheles roettenbacheri (Munster, 1839), without catalogue number (private collection), ultraviolet illumina-

tion (x 2); 2) Palaeopentacheles roetienbacheri (Munster, 1839), without catalogue number (BSPG), detail of thè chela of pereiopod

I (x 4); 3) Palaeopentacheles roettenbacheri (Munster, 1839), without catalogue number (BSPG), detail of abdominal somites (x 4);

4) Palaeopentacheles roettenbacheri (Munster, 1839), without catalogue number (BSPG), detail of thè tail fan (x 2 5)- 5) Palinurina

longipes Munster, 1839, lectotype, n. cat. AS V 46 (BSPG) (x 0.7).

THE UPPER JURASSIC SOLNHOFEN DECAPOD CRUSTACEAN FAUNA. PART I

63

PI. IXX - 1) Palinurina pygmea Miinster, 1839, lectotype, n. cat. AS I 996 (BSPG) (x 1.5); 2) Palinurina tenera Oppel, 1862, lecto-

type, n. cat. AS I 995 (BSPG) (x 1.2); 3) Palinurina tenera Oppel, 1862, n. cat. SMNS 65476 (Frattigiani collection) (naturai size); 4)

Cancrinos claviger Miinster, 1839, neotype, n. cat. AS I 998 (BSPG) (x 0.6).

64

ALESSANDRO GARASSINO & GUNTER SCHWEIGERT

PI. XX - 1) Cancrinos claviger Miinster, 1839, neotype, n. cat. AS I 998 (BSPG), detail of thè carapace and cephalic appendages

(naturai size); 2) Cancrinos claviger Miinster, 1839, neotype, n. cat. AS I 998 (BSPG), detail of thè antennal flagellum (x 1.7); 3)

( ancrinos latipes Mùnster, 1839, holotype by monotypy, n. cat. AS VII 769 (BSPG) (x 0.6); 4) Cancrinos claviger Miinster, 1839, n.

cat. 1975 IX 13 (BSPG) (x 0.5); 5) Cancrinos claviger Miinster, 1839, n. cat. 1975 IX 13 (BSPG), detail of thè pleurae of abdominal

somites and tail fan (x 1.2).

Ili - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Al¬

pe Turati (Erba, Como). Generi Hildoceras, Phymatoceras, Paronice-

ras e Frech iella. Conclusioni generali, pp. 143-204, 2 figg.. 6 tavv.

Volume XVIII

I - PINNA G., 1969 - Revisione delle ammoniti figurate da Giuseppe Me¬

neghini nelle Taw. 1-22 della « Monographie des fossiles du calcaire

rouge ammonitique » (1867-1881)./?/). 5-22, 2 figg., 6 tavv.

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

d’Orta ).pp. 23-92, 42 figg.. 4 taw. n.t.

Ili - PETRUCC1 F., BORTOLAM1 G. C. & DAL PIAZ G. V., 1970 - Ri¬

cerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul

suo substrato cristallino./)/). 93-169, con carta a colori al 1:40.000, 14

figg., 4 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./)/). 5-46, 2 tabb. n.t.

II - PINNA G. & LEVI-SETTI F., 1971 - 1 Dactylioceratidae della Provin¬

cia Mediterranea (Cephalopoda Ammonoidea). pp. 47-136, 21 figg.,

12 taw

III - PELOSIO G., 1973 - Le ammoniti del Trias medio di Asklepieion (Ar-

golide, Grecia) - 1. Fauna del «calcare a Ptychites» (Anisico sup.). pp.

137-168, 3 figg., 9 taw.

Volume XX

I - CORNAGGIA CASTIGLIONI 0., 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,

3 7 figg., 6 carte tematiche.

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

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

fflr

Volume XXI

I - PINNA G., 1 974 - 1 crostacei della fauna triassica di Cene in Val Seriana

(Bergamo)./)/). 5-34, 16 figg., 16 taw.

II - POLUZZI A., 1975 - I Briozoi Cheilostomi del Pliocene della Val d’Ar-

da (Piacenza, Italia)./)/). 35-78, 6 figg., 5 taw.

Ili - BRAMBILLA G., 1976 - I Molluschi pliocenici di Villalvemia (Ales¬

sandria). I. Lamellibranchi. pp. 79-128, 4 figg., 10 taw.

Volume XXII

I - CORNAGGIA CASTIGLIONI O. & CALEGARI G„ 1978 - Corpus

delle pintaderas preistoriche italiane. Problematica, schede, iconogra¬

fia. pp. 5-30, 6 figg., 13 tavv.

II - PINNA G., 1979 - Osteologia dello scheletro di Kritosaurus notabilis

(Lambe, 1914) del Museo Civico di Storia Naturale di Milano (Ornithi-

schia Hadrosauridae). pp. 31-56, 3 figg., 9 tavv.

Ili - BIANCOTTI A., 1981 - Geomorfologia dell’Alta Langa (Piemonte

meridionale). pp. 57-104, 28 figg., 12 tabb., 1 carta f.t.

Volume XXIII

I - GIACOBINI G., CALEGARI G. & PINNA G., 1982-1 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 figg., 16 tavv.

II - POLUZZI A., 1982 - I Radiolari quaternari di un ambiente idrotermale

del Mar Tirreno./)/). 45-72, 3 figg., 1 tab., 13 tavv.

Ili - ROSSI F., 1984 - Ammoniti del Kimmeridgiano superiore-Berriasiano

inferiore del Passo del Furio (Appennino Umbro-Marchigiano), pp. 73-

138, 9 figg.. 2 tabb., 8 taw.

Volume XXIV

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

sauro triassico del sottordine Lacertilia. pp. 5-28, 12 figg., 2 tavv.

II - NOSOTTI S. e PINNA G., 1989 - Storia delle ricerche e degli studi

sui rettili Placodonti. Parte prima 1830-1902 . pp. 29-86, 24 figg.. 12

taw.

Volume XXV

I - CALEGARI G., 1989 - Le incisioni rupestri di Taouardei (Gao, Mali).

Problematica generale e repertorio iconografico, pp. 1-14, 9 figg., 24

taw.

II - PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia funzionale

e paleoecologia del rettile placodonte Psephoderma alpinum Meyer,

1858 .pp. 15-50, 20 figg., 9 taw.

HI - CALDARA R., 1990 - Revisione Tassonomica delle specie paleartiche

del genere Tychius Germar (Coleoptera Curculionidae). pp. 51-218,

57 5 figg

Volume XXVI

I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia, Placo-

dontia ).pp. 1-21, 23 figg.

II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara preisto¬

rico: dati e interpretazioni./)/). 25-556, 647 figg.

Ili - ANDRI E. e ROSSI F., 1993 - Genesi ed evoluzione di frangenti,

cinture, barriere ed atolli. Dalle stromatoliti alle comunità di scogliera

moderne./)/). 559-610, 49 figg., 1 tav.

Volume XXVII

I - PINNA G. and GHISELIN M. edited by, 1996 - Biology as History. N.

1 . Systematic Biology as an Historical Science, pp. 1-133, 68 figs.

II - LEONARDI C. e SASSI D. a cura di, 1997 - Studi geobotanici ed en-

tomofaunistici nel Parco Regionale del Monte Barro, pp. 135-266, 122

figg -, 23 tabb.

Volume XXVIII

I - BANFI E. & GALASSO G., 1998 - La flora spontanea della città di

Milano alle soglie del terzo millennio e i suoi cambiamenti a partire dal

1700 .pp. 267-388, 71 figg., 30 tabb.

Volume XXIX

I - CALEGARI G., 1 999 - L’arte rupestre dell’Eritrea. Repertorio ragionato

ed esegesi iconografica./)/). 1-174, 268 figg.

Volume XXX

I - PEZZOTTA F. edited by, 2000 - Mineralogy and petrology of shallow

depth pegmatites. Paper from thè First International Workshop, pp. 1-

117, 30 figs., 19 tabs.

II - PARISI B„ FRANCHINO A. & BERTI A. con la collaborazione di

POTENZA B. & RUBINI D., 2000 - La Società Italiana di Scienze

Naturali 1855 - 2000. Percorsi storici./)/). 1-163, 199 figg.

III - DE ANGELI A. & GARASSINO A., 2002 - Galatheid, chirostylid and

porcellanid decapods (Crostacea, Decapoda, Anomura) from thè Eoce¬

ne and Oligocene of Vicenza (N Italy). /)/). 1-31, 21 figs., 9 pls.

Volume XXXI

I - NOSOTTI S. & RIEPPEL O., 2002 - The braincase of Placodus Agassiz,

1833 (Reptilia, Placodontia). pp. 1-18, 15 figs.

II - MARTORELLI G., 2002 - Monografia illustrata degli uccelli di rapina

in Italia. (1895). Riedizione a cura di Fausto Barbagli, pp. [XX] 1-216,

[14] 46 figg., 4 taw.

III - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphaigis dalsassoi n. gen.

n. sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano

(Lombardy, N Italy)./)/). 1-33, 19 figs., 1 tab., 3 pls.

Volume XXXII

I - ALESSANDRELLO A., BRACCHI G. & RIOU B„ 2004 - Polychaete,

sipunculan and enteropneust worms from thè Lower Callovian (Middle

Jurassic) of La Voulte-sur-Rhóne (Ardèche, France). pp. 1-16, 9 figs.,

1 pi¬

li - RIEPPEL O. & HEAD J. J., 2004 - New specimens of thè fossil snake

genus Eupodophis Rage & Escuillié, from Cenomanian (Late Creta-

ceous) of Lebanon. pp. 1-26, 13 figs., 1 tab.

III - BRACCHI G. & ALESSANDRELLO A., 2005 - Paleodiversity of thè

free-living polychaetes (Annelida, Polychaeta) and description of new

taxa from thè Upper Cretaceous Lagerstdtten of Haqel, Hadjula and

Al-Namoura (Lebanon). pp. 1-48, 8 figs., 1 tab., 16 pls.

Volume XXXIII

I - BOESI A. & CARDI F. edited by, 2005 - Wildlife and plants in tradi-

tional and modem Tibet: conception, exploration and conservation. pp.

1-88, 30 figs., 9 tabs.

II - BANFI E., BRACCHI G„ GALASSO . & ROMANI E., 2005 - Agro-

stologia Piacentina, pp. , figs., tabs.

III - LIVI P. a cura di 2005 - I fondi speciali della Biblioteca del Museo

Civico di Storia Naturale di Milano. La raccolta di stampe antiche del

Centro Studi Archeologia Africana, pp. 1-250, 389 figs.

Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali,

Museo Civico di Storia Naturale, Corso Venezia 55 -20121 Milano

Pubblicazione disponibile al cambio

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Volume XXXIV - Fascicolo II

della Società Italiana

di Scienze Naturali

e del Museo Civico

di Storia Naturale di Milano

DIRK FUCHS

MORPHOLOGY, TAXONOMY AND DIVERSITY

OF VAMPYROPOD COLEOIDS

(CEPHALOPODA)

FROM THE UPPER CRETACEOUS

OF LEBANON

MILANO MAGGIO 2006

Elenco delle Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

Volume I

I - CORNALI A E., 1865 - Descrizione di una nuova specie dei genere

Felis: Felis jacobita (Com.). 9 pp., 1 tav.

II - MAGNI-GRIFF1 F., 1865 - Di una specie d 'Hippolais nuova per l’Ita¬

lia. 6 pp., 1 tav.

Ili - GASTALDI B., 1865 - Sulla riescavazione dei bacini lacustri per opera

degli antichi ghiacciai. 30 pp., 2 figg., 2 tavv.

IV - SEGUENZA G., 1865 - Paleontologia malacologica dei terreni terziarii

del distretto di Messina. 88 pp., 8 taw.

V - GIBELLI G., 1865 - Sugli organi riproduttori del genere Verrucaria,

16 pp., 1 tav.

VI - BEGGIATO F. S., 1865 - Antracoterio di Zovencedo e di Monteviale

nel Vicentino, 10 pp., 1 tav.

VII - COCCHI I., 1865 - Di alcuni resti umani e degli oggetti di umana

industria dei tempi preistorici raccolti in Toscana. 32 pp., 4 taw.

Vili - TARGIONI-TOZZETTI A., 1866 - Come sia fatto l’organo che fa

lume nella lucciola volante dell’Italia centrale (Luciola italica) e come

le fibre muscolari in questo ed altri Insetti ed Artropodi. 28 pp., 2 taw.

IX - MAGGI L., 1865 - Intorno al genere Aeolosoma. 18 pp., 2 tavv.

X - CORNALIA E., 1 865 - Sopra i caratteri microscopici offerti dalle Cantari¬

di e da altri Coleotteri facili a confondersi con esse. 40 pp., 4 taw.

Volume II

I - ISSEL A., 1866 - Dei Molluschi raccolti nella provincia di Pisa, 38 pp.

II - GENTILLI A., 1 866 - Quelques considérations sur l’origine des bassins

lacustres, àpropos des sondages du Lac de Come. 12 pp., 8 ta\>v.

Ili - MOLON F., 1 867 - Sulla flora terziaria delle Prealpi venete. 140 pp.

IV - D’ACHIARDI A., 1866 - Corallarj fossili del terreno nummuiitico

delle Alpi venete. 54 pp., 5 taw.

V - COCCHI I., 1 866 - 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.

VII - COCCHI I., 1 866 - L’uomo fossile nell’Italia centrale. 82 pp., 21 figg.,

4 taw.

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 Vìpera Redii. 16 pp., 1 tav.

II - GARO VAGLIO S., 1 867 - Thelopsis, Belonia, Weitenwebera et Limbo¬

ria, quatuor Lichenum Angiocarporum genera recognita iconibusque

illustrata. 12 pp., 2 taw.

III - TARGIONI-TOZZETTI A., 1867 - Studii sulle Cocciniglie. 88 pp., 7 taw.

IV - CLAPAREDE 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 taw.

Volume IV

I - D'ACHIARDI A., 1868 - Corallarj fossili del terreno nummuiitico del-

l’Alpi venete. Parte 11. 32 pp. 8 tavv.

II - GARO VAGLIO S., 1868 - Octona Lichenum genera vel adirne con¬

troversa, ve 1 sedis prorsus incertae in systemate, novis descriptionibus

iconibusque accuratissimis illustrata. 18 pp., 2 taw.

Ili - MARINONI C., 1868 - Le abitazioni lacustri e gli avanzi di umana

industria in Lombardia. 66 pp., 5 figg., 7 taw.

IV - (Non pubblicato).

V - MARINONI C., 1871 - Nuovi avanzi preistorici in Lombardia. 28 pp.,

3 figg., 2 taw.

NUOVA SERIE

Volume V

I - MARTORELLI G., 1895 - Monografia illustrata degli uccelli di rapina

in Italia. 216 pp., 46 figg., 4 taw.

Volume VI

I - DE ALESSANDRI G., 1897 - La pietra da cantoni di Rosignano e di

Vignale. Studi stratigrafici e paleontologici. 104 pp., 2 taw., 1 carta.

II - MARIOREL.LI 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 taw.

Volume Vili

1 - REPOSSI E., 1915 - La bassa Valle della Mera. Studi petrografici e

geologici. Parte 1 .pp. 1-46, 5 figg., 3 taw.

II - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi petrogra¬

fici e geologici. Parte II . pp. 47-186, 5 figg. 9 taw.

Ili - A1RAGHI C., 1917 - Sui molari d’elefante delle alluvioni lombarde,

con osservazioni sulla filogenia e scomparsa di alcuni Proboscidati./?/?.

187-242, 4 figg., 3 taw.

Volume IX

I - BEZZI M., 1918 - Studi sulla ditterofauna nivale delle Alpi italiane./?/?.

1-164. 7 figg. 2 taw.

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 tavv.

Volume X

I - DESIO A., 1929 - Studi geologici sulla regione dell’Albenza (Prealpi

Bergamasche)./?/?. 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- I recettori degli Agamidi. pp. 209-326, 80 figg.

Volume XI

I - GUIGL1A D., 1 944 - Gli Sfecidi italiani del Museo di Milano (Hymen.).

pp. 1-44, 4 figg., 5 tavv.

II-III - GIACOBINI V. e P1GNATTI 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)./?/?. 1-70, 4 figg. 6 tavv.

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./?/?. 1-64, 25 figg., 9 taw., 1 carta.

II - PINNA G., 1963 - Ammoniti del Lias superiore (Toarciano) dell’Alpe

Turati (Erba, Como). Generi Mercaticeras, Pseudomercaticeras e Bro-

dieìa. pp. 65-98, 2 figg., 4 tavv.

Ili - ZANZUCCHI G., 1963 - Le Ammoniti del Lias superiore (Toarciano)

di Entratico in Val Cavallina (Bergamasco orientale), pp. 99-146, 2

figg., 8 tavv.

Volume XIV

I - VENZO S., 1965 - Rilevamento geologico dell’anfiteatro morenico

frontale 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 tavv.

Ili - DI ENI I., MASSARI F. e MONTANARI L., 1966 - Il Paleogene dei

dintorni di Orosei (Sardegna), pp. 13-184, 5 figg., 8 taw.

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, 2 7 figg. 9 tavv.

II - DIENI I. e MASSARI F., 1966 - Il Neogene e il Quaternario dei dintorni

di Orosei (Sardegna)./?/?. 89-142, 8 figg., 7 taw.

Ili - BARBIERI F„ IACCARINO S„ BARBIERI F. & PETRUCCI F„ 1967

- Il Pliocene del Subappennino Piacentino-Parmense-Reggiano. pp.

143-188, 20 figg., 3 tavv.

Volume XVI

I - CARETTO P. G., 1967 - Studio morfologico con l’ausilio del metodo

statistico e nuova classificazione dei Gasteropodi pliocenici attribuibili

al Murex brandaris Linneo, pp. 1-60, 1 fig., 7 tabb., 10 taw.

II - SACCHI VIALLI G. e CANTALUPPI G„ 1967-1 nuovi fossili di Goz¬

zano (Prealpi piemontesi)./?/?. 61-128, 30 figg., 8 taw.

Ili - PIGORINI B., 1967 - Aspetti sedimentologici del Mare Adriatico, pp.

129-200, 13 figg., 4 tabb. 7 tavv.

Volume XVII

I - PINNA G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Alpe

Turati (Erba, Como). Famiglie Lytoceratidae, Nannolytoceratidae,

Hammatoceratidae (excl. Phymatoceratinae) Hildoceratidae (excl.

Hildoceratinae e Bouleiceratinae) . pp. 1-70, 2 taw. n.t., 6 figg-, 6 taw.

II - VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi del-

1 Anisico superiore di Lenna in Val Brembana (Bergamo)./?/?. 71-142,

5 figg., 11 taw.

Dirk Fuchs

Freie Universitat Berlin, Institut fìir Geologische Wissenschaften, Fachrichtung Palàontologie

Morphology, taxonomy and diversity

of vampyropod Coleoids (Cephalopoda)

from thè Upper Cretaceous of Lebanon

Volume XXXIV - Fascicolo II

Maggio 2006

Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

© 2006 Società Italiana di Scienze Naturali

Museo Civico di Storia Naturale di Milano

Corso Venezia , 55 - 20121 Milano

In copertina: Vampyroteuthis infernalìs (Chun, 1903). Drawing by Monika Bulang-Lòrcher.

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 - Maggio 2006

ISSN 0376-2726

Dirk Fuchs

Morphology, taxonomy and diversity

of vampyropod Coleoids (Cephalopoda)

from thè Upper Cretaceous of Lebanon

Abstract - Morphology, taxonomy and diversity of coleoids from thè Cenomanian and Santonian of Lebanon are described. Until

hitherto, thè coleoid fauna of thè Late Cretaceous Lithograhic Limestones of Hàqel, Hjoùla (both Upper Cenomanian) and Sàhel Aalma

(Upper Santonian) includes five vampyropod and only one belemnoid species. Decabrachia are unknown from these localities.

The comparatively poor information about thè genus Dorateuthis Woodward, 1883 is discussed morphologically as well as

taxonomically. Rachiteuthis donovani, a taxon newly introduced herein, is described for thè first time. Specimens determined as Tra-

chyteuthis cf. hastiformis Riippel, 1 829 extend thè stratigraphical range of this genus from thè Callovian up to thè Cenomanian. Close

morphological affinities of Libanoteuthis libanotica Fraas, 1878 with Glyphiteuthis ornata Reuss, 1854 from thè Turonian of Bohemia

(Czech Republic) suggest that Libanoteuthis Kretzoi, 1942 is a junior subjective synonym of Glyphiteuthis Reuss, 1854. Morphologi¬

cal knowledge about each taxon is improved by an exceptional high number of specimens, which are deposited in thè Museo Civico

di Storia Naturale di Milano (MSNM).

Key-words: Coleoidea, Vampyropoda, Cenomanian, Santonian, Lebanon.

Riassunto - Morfologia, tassonomia e diversità dei coleoidei vampiropodi (Cephalopoda) del Cretaceo superiore del Libano.

Vengono descritte la morfologia, la tassonomia e la diversità dei coleoidei del Cenomaniano e del Santoniano del Libano. Ad oggi,

la fauna a coleoidei dei calcari litografici del Cretaceo superiore di Hàqel, di Hijoula (entrambi Cenomaniano superiore) e di Sàhel

Aalma (Santoniano superiore) comprende cinque specie di vampiropodi e una sola specie di belemnoidi. In queste località non sono

stati rinvenuti esemplari di Decabrachia.

Vengono discusse dal punto di vista morfologico e tassonomico le informazioni relativamente scarse sul genere Dorateuthis

Woodward, 1883. Viene descritto per la prima volta Rachiteuthis donovani, un taxon precedentemente istituito e qui ripresentato. Gli

esemplari ascritti a Trachyteuthis cf. hastiformis Riippel, 1829 estendono la distribuzione stratigrafica di questo genere dal Calloviano

fino al Cenomaniano. Le strette affinità morfologiche fra Libanoteuthis libanotica Fraas, 1878 e Glyphiteuthis ornata Reuss, 1854 del

Turoniano della Boemia (Repubblica Ceca) inducono a considerare Libanoteuthis Kretzoi, 1942 come sinonimo juniore soggettivo di

Glyphiteuthis Reuss, 1 854. La conoscenza della morfologia di ogni taxon è accresciuta grazie al numero eccezionalmente alto degli

esemplari a disposizione, depositati presso il Museo Civico di Storia Naturale di Milano (MSNM).

Parole chiave: Coleoidea, Vampyropoda, Cenomaniano, Santoniano, Libano.

INTRODUCTION

The Lithographic Limestones (“fish beds”, “fish

shales”) of Lebanon (Hàqel, Hjoùla and Sàhel Aalma) are

well-known for their well-preserved fossil assemblage.

Especially fishes and crustaceans have been thè subject of

many publications (Garassino, 1994; Schram et al., 1999;

Forey et al., 2003; Taverne & Filleul, 2003). By contrast,

knowledge about Cephalopod coleoids is comparatively

poor, although many beautiful specimens occur.

Coleoidea are typifìed by thè extant squids, cuttlefishes

and octopuses. According to Boletzky (2003) recent Coleoi¬

dea are classified into thè eight-armed Vampyropoda and thè

ten-armed Decabrachia. The third coleoid superorder consti-

tutes thè extinct Belemnoidea (Late Carboniferous - Late

Cretaceous). Within thè Vampyropoda thè Cirroctopoda and

thè Octopoda are grouped together as Octobrachia. The Octo-

brachia represent thè sister taxon of thè Vampyromorpha.

Decabrachia include thè Sepiida, Spirulida, Idi-

osepiida, Sepiolida and thè Teuthida. Although these

groups are comparatively well known, their phylogenetic

relationships are surprisingly problematic (Young & Vec¬

chione, 1996; Vecchione et al., 2000; Boletzky, 2003).

The fossil record of coleoid groups is usually biased

towards thè preservation of mineralised “hard parts”.

Discoveries of soft tissues are exclusively connected to

so-called “Konservat-Lagerstàtten”. Their extraordinary

preservational potential is thè most important factor by

which thè poor morphological knowledge, and conse-

quently thè problematic phylogeny of thè Coleoidea can

be improved (Donovan, 1977; Engeser, 1990; Doyle et

al., 1994; Young et al., 1998; Boletzky, 1999; Haas, 2002

and 2003). Well-known Lagerstàtten with a comparatively

high soft-part resolution are, for example, thè Sinemurian

of Osteno (Italy), thè Toarcian of Holzmaden (Germany)

and Yorkshire (England), thè Callovian of La Voulte-

sur-Rhòne (France) and Christian Malford (England),

thè Tithonian of Solnhofen (Germany), thè Cenomanian

of Hàqel (Lebanon) and thè Santonian of Sàhel Aalma

(Lebanon). For palaeobiologists, these biotas form unique

taphonomic and evolutionary Windows through thè Meso-

zoic, allowing detailed reconstructions.

DIRK FUCHS

The Late Cretaceous Lithographic Limestones of

Lebanon display a soft-part preservation that rivals thè

famous German “Posidonienschiefer” of Holzmaden and

thè “Solnhofener Plattenkalke”. So far five coleoid species

{Dorateuthis syriaca, Dorateuthis sahilalmae, Libano-

teuthis libanotica, Palaeoctopus newboldi, Acanthoteuthis

syriaca ) are known from these Late Cretaceous deposits.

Several specimens of them (including Holotypes) are

deposited in thè museums of Berlin, Stuttgart (both Ger-

many), Paris (France) and London (United Kingdom).

However, thè Museo Civico di Storia Naturale di Milano

(Italy) as well as thè Black Hills Institute of Geological

Research in Black Hills (United States) house by far thè j

largest collection of Lebanese coleoids. Neither thè Milan j

nor thè Black Hills specimens have ever been described.

For thè first time a large number of specimens allows a

detailed morphological documentation and comparison.

The description of both collections starts with thè Milan

specimens. A forthcoming project with Neal L. Larson

(Black Hills Institute of Geological Research) describing

thè specimens of thè Black Hills collection will comprise I

thè entire cephalopod fauna of Hàqel.

PREVIOUS STUDIES ON COLEOID CEPHALOPODS FROM THE LATE CRETACEOUS

OF LEBANON

First investigations of Lebanese coleoids began in

1846 with incomplete descriptions by Sowerby, who first

mentioned Calai's Newboldi, better known as Palaeoc¬

topus newboldi Woodward, 1883, thè first undoubted

member of thè Octopoda. More serious and detailed

studies by Lartret (1877), Fraas (1878), and Woodward

(1883, 1896) yielded two other members of thè Cole¬

oidea which are now assigned to thè Vampyropoda:

Dorateuthis syriaca Woodward, 1883 and Libanoteuthis

libanotica Fraas, 1878.

In 1 922 Naef established two additional taxa. One was

erected by means of a doubtful specimen of Fraas, which

Naef called “ Sepìalites Sahil Almae ”. This specimen

(deposited in Stuttgart) is now considered to be thè Holo-

type of Dorateuthis sahilalmae. Naef’s other specimen,

“ Beloteuthis libanotica ”, was proved to be thè isolated

gladius element of Palaeoctopus newboldi (Haas, 2002;

Bizikov, 2004).

Roger ( 1 946) treated taxa such as Palaeoctopus new¬

boldi, Dorateuthis syriaca and Libanoteuthis libanotica

and established soft parts and hooks of thè first Lebanese

belemnoid Belemnoteuthis (Acanthoteuthis) syriaca,

whose classification is admittedly uncertain.

Additional information on Lebanese coleoids Comes

from Engeser & Reitner (1986). They re-described thè

collection of Fraas as well as other Lebanese specimens

accumulated in thè Museum fur Naturkunde Stuttgart

(SMNS). They erected a new genus ( Parateudopsis ) and

mentioned an uncertain species ( Palaeololigo sp.). Both s

taxa have proved to belong to remains of Palaeoctopus.

GEOLOGICAL SETTING

Studied specimens are derived from thè Cenomanian

outcrops at Hàqel (45 km north of Beirut) and Hjoula (10

km east of Byblos) as well as thè Santonian outcrops of

Sàhel Aalma (25 km north of Beirut) (Fig. 1). Today each

site is exclusively quarried for fossils. In recent decades

several authors have investigated these geological settings

(Ejel & Dubertret, 1966; Hùckel, 1970, 1974 and 1974b;

Hemleben, 1977; Dalla Vecchia et al., 2002).

Fig. 1 - Locality map of Hàqel, Hjoùla and Sàhel Aalma.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

5

The precise age of Hàqel and Hjoùla varies accord-

ing to different authors. In his comprehensive sedimen-

tological study Huckel (1970) divided thè Cenomanian

Limestones into seven units and concluded that Hàqel

(unite V) and Hjoùla (unite IV) were deposited during thè

later Lower Cenomanian. Owing to thè occurrence of thè

ammonite Allocrioceras annulatum as a member of thè

Metoicoceras geslinianum Zone, Wippich & Lehmann

(2004) confirmed thè conclusions of Hemleben ( 1 977) and

dated Hàqel and Hjoùla as Upper Cenomanian (Fig. 2).

Dalla Vecchia et al. (2002) considered Hàqel and Hjoùla

as lower Upper Cenomanian, because Nammoùra (another

fossil-rich locality a few kilometres in thè South of Hjoùla

and slightly younger than Hàqel and Hjoùla) is dated as

Middle Cenomanian

Sàhel Aalma, a locality with very similar depositional

conditions and almost thè same coleoid diversity, is com-

monly dated as Upper Santonian (Ejel-Dubertret, 1966;

Garassino, 1994).

Each outcrop is of particular morphological impor-

tance, because fossils show precise preservation of soft

tissues. The fauna consists of fishes (Forey et al, 2003;

Taverne & Filleul, 2003), crustaceans (Garassino, 1994;

Schram et al., 1999), echinoderms (Huckel, 1970), other

molluscs (Forey et al., 2003; Wippich & Lehmann, 2004)

and worms (Dalla Vecchia & Venturini, 1999; Bracchi &

Alessandrello, 2005), pterosaurs (Dalla Vecchia et al.,

2001) and other reptiles (Dal Sasso & Pinna, 1997; Dalla

Vecchia & Venturini 1999; Rieppel & Head, 2004; Cald-

well, 2006 in press; Tong et al., 2006).

Palaeogeographers believe that thè sediments of Hàqel

and Hjoùla were deposited at thè edge of thè Thethian

shelf margin, 400 km away from thè ancient coastline

(Hembleben, 1977; Philip et al., 1993). The map in Fig. 3

illustrates thè palaeogeographic situation of thè Central

Tethys during thè Cenomanian. Huckel (1970) found

that Hàqel and Hjoùla were deposited in small basins.

Most authors agree that thè “fish beds” were formed in

an anoxic environment. Hemleben (1977) suggested that

upwelling at thè shelf margin occasionally supported

blooms of protists or “red tides”, which caused episodes

of mass mortality. Deoxygenated bottom water and high

sedimentation rates then prevented thè decay of dead

bodies.

< si

o

w

u

s

w

QZ

U

pts

Ed

Cd

Cu

D

84mya

86mya .

89mya •

94mya

■ lOOmya ■

Campanian

Santonian

Coniacian

Turonian

Cenomanian

Albian

M

Sahel Aalma

Hàqel

Hjoula

Fig. 2 - Stratigraphic age of Hàqel, Hjoùla and Sàhel Aalma.

Russian Platform

cxposed land

HH

shallow marine

deep marin

North- Africa

Fennoscandia

Fig. 3 - Palaeogeographic map of thè Central Tethys during thè Late

Cenomanian. L = Lebanon. Modified after Philip et al. (1993).

MATERIAL

The study was conducted on 38 specimens belong-

ing to thè Museo Civico di Storia Naturale di Milano

(MSNM). Most specimens come from Hàqel and Hjoùla.

Only two specimens come from Sàhel Aalma. Material

from Nammoùra was not available.

Specimens were investigated in detail with a binocular

microscope. Oblique light reveals faint structures, which

were not recognized under vertical light conditions. Ultra-

violet light was used to indicate phosphatised soft tissues.

A number of additional specimens from different locali-

ties (including Holotypes housed in collections of Berlin,

Stuttgart, London, Paris Vienna, and Prague) were used as

comparative material.

MODES OF PRESERVATION

All specimens are compressed and flattened on a 2-

dimensional surface. Although thè mineralogy was not

analysed, gladii (singular “gladius”: a stiff supporting

structure in thè dorsal mantle, see below) were most

probably preserved as francolite. However, francolite is

a diagenetical result. On thè assumption that vampyropod

gladii derived from a belemnoid organic proostracum,

a purely organic composition is reasonable (Hewitt &

Wignall, 1988; Doguzhaeva & Mutvei, 2003; Fuchs et

al., submitted).

6

DIRK FUCHS

Konservat-Lagerstàtten are characterised by a detailed

preservation of soft tissues. According to Allison (1988),

Allison & Briggs (1991), Wilby (1993), Kear et al. (1995)

and Wilby & Briggs (1997), thè highest-fidelity preserva¬

tion occurs where muscular structures are post-mortem

mineralised in apatite (a calciumphosphate).

Sofìt-part preservation of coleoids is various. Only

in rare cases thè specimens include articulated bodies

complete with mantle, arms, retractors and fins. Typically,

coleoids are only partially mineralised and disarticulated,

but mineralisation of thè mantle in dose association with

ami stubs occurs often. Jaw preservation is common, and

occasionally stomach contents as well.

An ink sac occurs in almost each specimen, but thè

possession of ink is not considered to be a diagnostic

feature within coleoids and is therefore neglected in thè

descriptions.

SYSTEMATIC PALAEONTOLOGY

Subclass Coleoidea Bather, 1888

Superorder Vampyropoda Boletzky, 1992

Suborder Prototeuthina Naef, 1921

Family Plesioteuthididae Naef, 1921

Genus Dorateuthis Woodward, 1883

Type species: Dorateuthis Syriaca Woodward, 1883;

by monotypy.

Stratigraphical range: Upper Cenomanian of Hjoùla

and Upper Santonian of Sàhel, Lebanon.

Remarks: Woodward (1883) first described Dora¬

teuthis syriaca from Sàhel Aalma. Later-treated specimens

of Dorateuthis syriaca as well as specimens assigned to

Dorateuthis sahilalmae derived from Sàhel Aalma. How-

ever, as it will be shown, Dorateuthis already occurred in

thè older Cenomanian of Hjoùla.

The placement of thè genus Dorateuthis within thè

Leptotheuthididae, Naef 1922 by Jeletzky (1966: 45) and

other authors cannot be accepted. The view of Jeletzky

(1966) is based on a doubtful reconstruction given by

Roger (1946: 14, Figs. 6-7). After re-study of Roger’s

specimens in Paris there are no doubts that thè genus

Dorateuthis has more in common with Plesioteuthis

or Senefelderiteuthis than with Leptotheuthis. Thus we

follow thè classification of Donovan (1977) and Engeser

& Reitner (1986) in placing thè genus Dorateuthis within

thè Plesioteuthididae.

Dorateuthis cf. syriaca Woodward, 1883 (Pls. I-III)

1 883 Dorateuthis Syriaca n. sp. - Woodward, p. 1 ff., PI. 1 .

1888 Curculionites senonicus - Kolbe, pp. 135-136, PI.

11, Fig. 8.

1920 Dorateuthis Syriaca Woodward - Bùlow-Trummer,

p. 262.

1922 Sepialites Sahil-Almae - Naef, p. 134, Fig. 49.

1922 Dorateuthis syriaca Woodward, 1883 - Naef, p.

135, Fig. 50.

(?) 1 932 Doratoteuthis syriaca H. Woodward - Quenstedt,

p. 186 ff.

1946 Leptoteuthis syriaca Woodward - Roger, p. 14 ff.,

Fig. 6-7, PI. 4, Fig. 5-6, PI. 9, Fig. 1.

1966 Dorateuthis syriaca - Jeletzky, pp. 44-45.

1977 Plesioteuthis syriaca - Donovan, p. 40.

1982 Dorateuthis syriaca Woodward - Reitner & Enge¬

ser, p. 215.

1986 Dorateuthis syriaca Woodward, 1883 - Engeser &

Reitner, p. 4.

1986 Dorateuthis sahilalmae (Naef, 1922) - Engeser &

Reitner, p. 4, Fig 1, PI. 1, Fig. 1.

1 986 Dorateuthis sp. - Engeser & Reitner, p. 5, PI. 1 , Fig. 2.

1987 Dorateuthis .syr/aca Woodward, 1883-Riegraf,p. 97.

1988 Dorateuthis syriaca Woodward, 1883 - Engeser, p. 42.

2002 Maioteuthis n. sp. - Novati 2002, p. 88.

2004 Dorateuthis syriaca Woodward — Lukeneder &

Harzhauser, p. 3.

Holotype: specimen of Woodward (1883, PI. 1), BMNH

C 5017 (PI. Ili, Fig. B).

Locus typicus: Sàhel Aalma, Lebanon.

Stratum typicum: Lithographic Limestones of Sàhel

Aalma, Upper Santonian.

Geographic and stratigraphic distribution: Upper

Cenomanian of Hjoùla - Upper Santonian of Sàhel

Aalma, Lebanon.

Material: eight specimens were investigated: MSNM

Ì25144 (PI. I, Figs. A-B), Ì24800 (PI. II, Fig. A), Ì25133

(PI. II, Fig. B), Ì25134 (PI. Ili, Fig. A), Ì25128, Ì24802

originated from Hjoùla, whereas i23 1 08 (PI. I, Fig. C) and

Ì26092 come from Sàhel Aalma.

Description

Gladius: specimens MSNM i25 144 (PI. I, Fig. A),

Ì23108 (PI. I, Fig. C), Ì25128 and Ì26092 are seen either

in dorsal or ventral views whereas specimens Ì24800 (PI.

II, Fig. A), Ì25133 (PI. II, Fig. B), Ì25134 (PI. Ili, Fig.

A) and Ì24802 are embedded laterally. Lateral specimens

provide less information about gladius characteristics. In

comparison to dorsoventral specimens, lateral examples

provide more information about soft-body morphology

than gladius characteristics. The reconstruction in Fig. 4

is only tentative, because each gladius of Dorateuthis is

in a bad condition.

Only in specimen MSNM i25 144 are imprints of a

complete gladius preserved (PI. I, Fig. A). It is embedded

dorsoventrally and seen in dorsal view. The total length of

thè gladius is 7.5 cm. Its anterior end is 0.8 cm in width.

Compared to thè length of other specimens MSNM

Ì25144 is a juvenile (MSNM i25 1 34 is longest with at

least 24 cm). More information about thè maximum width

of other specimens is not worthwhile owing to incomplete

preservation, compaction or lateral embedment.

In MSNM Ì25144 (PI. I, Fig. A) and Ì23108 (PI. I, Fig.

C) pronounced lateral ridges are rounded and ca. 1 mm in

width. Their increase in thickness throughout thè whole

length is insignificant. Both ridges diverge at an angle

of approximately 7°. Lateral ridges must be regarded as

lateral swellings of thè median field.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

7

CO?

lateral dorsal

view view

Fig. 4 - Dorateuthis syriaca. Reconstruction of thè gladius. The mor-

phology of thè posterior gladius end is uncertain. co = conus; lf = lateral

field; Lr = lateral ridge; mf = median field; mr = median ridge.

A distinct median line or ridge can be observed in

specimen MSNM i23 1 08 (PI. I, Fig. C) and Ì26092. In

thè latter thè line is seen to reach thè most anterior end of

thè gladius.

The anteriormost extremity of thè gladius and growth

lines on thè gladius surface are poorly preserved, but there

are no indications for a pointed anterior end.

Imprints on both sides of this posterior end observable

in specimen MSNM i25 144 (PI. I, Fig. B) are probably

imprints of lateral fields (conus fields). If they are indeed

lateral fields they are extremely narrow and short (2 mm

in width and 2.1 cm in length respectively). However, lat¬

eral specimens do not confimi this assumption.

Lateral specimens MSNM Ì24800 (PI. II, Fig. A) and

Ì25133 (PI. II, Fig. B) demonstrate thè originai curvature

of thè gladius, which probably reflects its tension within

thè mantle sac.

Arms: specimens MSNM i25 133, i25 1 34 and Ì26092

exhibit arm preservation. The first two show at least three

arms (PI. II, Fig. B; PI. Ili, Fig. A), thè latter at least four.

In this specimen arms preserved as phosphatised mus-

culature are only fragmentary and apparently tangled. It

seems worthwhile to note that thè diameters of thè arms

are relatively thick. In MSNM Ì26092 one arm is at least

5 cm long. Ultraviolet light supplies no further informa-

tion, so that reliable measuring data are not available.

Neither tentacles, clubs, suckers nor cirri are discemible

anywhere.

Head: in four specimens thè position of thè head can

be presumed (MSNM i25 133, i25 1 34, Ì24800, Ì26092)

from thè buccal mass. Strongly phosphatised structures

between thè arms and thè mantle interpreted as thè buccal

mass are displaced from thè mantle edge. In MSNM

Ì24800 (PI. II, Fig. A), Ì25133 (PI. II, Fig. B) and i25 1 34

(PI. Ili, Fig. A) thè buccal mass is located some distance

in front of thè anterior gladius end. Ultraviolet light

suggests that thè anterior end of thè gladius reaches thè

anterior mantle margin. This indicates that thè head was

apparently demarcated. Beaks were not observed.

Fins: in specimens MSNM i25 128 and i25 134 imprints

of fins are present. In lateral specimen MSNM i25 134 (PI.

Ili, Fig. A) preparation has laid open a ca. 3 cm long weak

lobate structure, which is considered to be a fin. Similar

lobate structures are also present in MSNM i25 128. How¬

ever, in both cases this is not beyond doubt, as ultraviolet

light did not confimi this assumption.

Stomach: in specimen MSNM i25 144 stomach contents

are preserved (PI. I, Fig. A). As in recent coleoids thè stom¬

ach is situated between thè ink sac and thè rear of thè mantle

cavity. Ingested food largely consists of fish remains, so

Dorateuthis was certainly an active predator. As correctly

suggested by Lukeneder & Harzhauser (2005) thè mass of

food indicates thè presence of a crop (caecum).

Remarks: thè Milan specimens of Dorateuthis syriaca

are thè first evidence of this taxon in thè Cenomanian.

However, morphological knowledge about thè genus

Dorateuthis (including both species) is stili comparatively

scarce. This is partly due to a very flimsy gladius. After

re-investigation of thè Holotypes of Dorateuthis syriaca

(PI. Ili, Fig. B) and Dorateuthis sahilalmae (PI. Ili, Fig.

C) and other examples of this genus in collections from

Berlin, Stuttgart, Vienna, Paris and London and compari-

son with thè Milan specimens described herein (including

thè Milan specimens a total amount of 27 specimens of

this genus were investigated) it can be concluded that

only thè lateral ridges and partly thè median ridge have

thè potential to withstand decay. Often we have just

hollow imprints of thè massive lateral ridges. Because of

its narrowness thè median field has apparently no preser-

vational potential. However, long and wide lateral fields

as suggested by Roger (1946) can be excluded. If lateral

fields were present in this genus, they must have been

extremely short and narrow as reconstmcted in Fig. 4.

Further, it cannot be ruled out that thè gladius is some-

times disrupted longitudinally. In thè author’s opinion,

differing information about apical angles ranging from

7° to 25° is a result of disruption and distortions. In

spite of this observation, thè two representatives of thè

genus are differentiated by means of their apical angles.

According to Engeser & Reitner (1986: 4) D. sahilalmae

(PI. Ili, Fig. C) is mainly distinguished from D. syriaca

by a larger apical angle and a bigger ink sac. The latter

is here considered to be highly dependent on thè volume

of ink within thè sac just before death and on random

preservational compaction. None of thè 27 specimens of

Dorateuthis investigated showed a correlation between

ink sac size and thè apical angle. Thus an unambiguous

character distinguishing D. sahilalmae from D. syriaca is

stili missing.

8

DIRK FUCHS

Re-investigation of thè arms of thè Holotypes of D.

syriaca Woodward, 1883 as well as D. sahilalmae Naef,

1922 has shown that this genus is certainly an eight-

armed coleoid. A specimen from thè Naturhistorisches

Museum Wien exhibits thè best-preserved arm crown

consisting of eight distinct arms (PI. II, Fig. C, originai

of Lukeneder & Harzhauser, 2004). There are absolutely

no indications for thè presence of tentacles as proposed

by Woodward (1883). However, in cases where arm mus-

culature is preserved it appears that some animals had

well-developed, fleshy arms (c.f. MSNM Ì26092), while

others seem to have had comparably thin arms (PI. II, Fig.

C; PI. Ili, Fig. B). A tentati ve ontogenetical correlation for

thickest and longest arms is observed in specimens with a

gladius longer than 20 cm. On thè other hand, thinner and

shorter arms are associated with gladii shorter than 10 cm.

Therefore different ontogenetical stages might explain

variable arm morphology. Altematively, inconsistencies

are more likely due to incorrect preparation or incomplete

preservation than to a specific or an ontogenetic charac-

ter. Unsuccessful correlation between arm thickness and

gladius characteristics confirms these considerations. As

a result, presence of two species within thè genus Dora-

teuthis remains doubtful.

Likewise, no distinct differences were found between

specimens from Hjoula, and Sàhel Aalma. Thus, gladius,

ink sac and arm morphology are stili useless as distin-

guishing characters. From Hàqel Dorateuthis is stili

unknown. Perhaps, thè undescribed material housed in

thè Black Hills Institute may provide additional informa-

tion. Until then, confirmation of two separated species

from Sàhel Aalma as well as erection of a new species

from Hjoula would be premature.

Suborder Teudopsidina Starobogatov, 1983

Family Palaeololiginidae Naef, 1921

Genus Rachiteuthis (n. gen.)

Derivatio nominis: from rachis-like appearance of thè

gladius.

Type species: Rachiteuthis donovani (n. sp., by mono-

typy)-

Locus typicus: Hjoula (Lebanon).

Stratum typicum: Lithographic Limestones of Hjoùla,

Upper Cenomanian.

Geographic and stratigraphic distribution: known only

from thè locus typicus and stratum typicum.

Diagnosis: palaeololiginid with a spear- or shaft-like

gladius; lateral fields spread out and short with a weak

hyperbolar zone; conus reduced; free rachis elongated;

distinct lateral ridges and a median unomamented keel

extend from posterior to anterior extremities; body tor-

pedo-shaped; insertions of triangular fins comparatively

broad; arms comparatively slender.

Remarks: thè genus is placed within thè Palaeololigi¬

nidae because of common possession of a sharply pointed

free rachis. Presence of lateral ridges similar to members

of thè Plesioteuthididae may also imply an affiliation to

these forms. However, in contrast to thè ventrally closed

true conus of Plesioteuthis, thè new genus shows a

reduced (opened) conus with outspread lateral fields.

Rachiteuthis donovani (n. sp.) (Pls. IV- VI)

1977 “PlesioteuthisA ike gladii” - Donovan, p. 40.

Derivatio nominis: thè name is in honour of D. T.

Donovan and his exceptional merits in palaeontological

malacology.

Holotype: MSNM Ì25142 (PI. IV).

Paratype I: MSNM i25 135 (PI. V, Fig. A).

Paratype II: MSNM Ì25139 (PI. V, Fig. B).

Locus typicus: Hjoùla (Lebanon).

Stratum typicum: Lithographic Limestones of Hjoùla,

Upper Cenomanian.

Geographic and stratigraphic distribution: known only

from thè locus typicus and stratum typicum.

Remarks: in 2002 Novati already recognized and sug-

gested this new taxon, but never described it. The taxon

name was therefore a nomen nudum. Present author

decided to keep thè taxon name and to present thè first

description of Rachiteuthis.

Material: five specimens were investigated: MSNM

Ì25142, Ì25139, Ì25135, i25132a+b and Ì25143. The

latter specimen is a juvenile and provides little additional

information, but thè former are adult and each of them

bears interesting features. Specimen MSNM i25 142 is

selected as thè Holotype because a complete gladius with

detailed structural morphology is preserved. Although thè

gladius of Paratype I (MSNM i25 135) is only preserved

as imprints, this specimen exhibits extraordinary soft-part

preservation, including arms, head, fiinnel, ink sac and

fins. Paratype II shows thè lateral view of this taxon.

Description

Gladius: thè Holotype (PI. IV) exhibits by far thè

best-preserved gladius. Because it is completely dorsov-

entrally embedded, thè obtained measurements are highly

usable (data from gladii of thè Paratypes are ambiguous

for they are more or less laterally compressed and are thus

not considered here). Additionally, gladius characteristics

of thè Holotype are preserved 3-dimensionally (Fig. 5).

Imprints of thè ventral surface are visible where gla¬

dius material is missing. The Holotypes gladius dimen-

sions (Fig. 5) are as following:

- gladius length (gl): 5.7 cm

- lateral field length (lfl): 1.3 cm

- median field width (mfwjf) at thè end of thè lateral fields:

0.35 cm

- maximum median field width (mfwmax): 0.59 cm

- maximum lateral field width (lfw): 0.82 cm

- lateral field length/gladius length (lfl/gl): 0.23

- median field width/gladius length (mfwmax /gl): 0.1

- median field width/gladius length (mfwjf /gl): 0.06

- lateral field width/lateral field length (lfw/lfl): 0.6

- apical angle (angle of diverging inner asymptotes): 8°

Lateral field length (lfl) accounts for 23% of total gla¬

dius length (gl). In other investigated specimens it varies

between 23 and 26%, but as already mentioned, this is pre-

sumably due to different degree of compaction. An unde-

termined specimen from Hjoùla in thè Museum of Naturai

History in London (Donovan, 1977: 40, “ plesioteuthis-like

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

M-mfw-H

Fig. 5 - Rachiteuthis donovani. Reconstruction of thè gladius. fri = free rachis length; gl = gladius length; ia = inner asymptote; lf =

lateral field; lfl = lateral field length; lfw = lateral field width; Ir = lateral ridge; mf = median field; mfw = median field width; mk =

median keel.

gladii”; register number BMNH C 28408) is identical with

thè present taxon and shows thè same lfl/gl - ratio of 0.23.

The maximum width of thè median field (mfwmax) is

0.59 cm, measured half way along thè rachis-like median

field (free rachis). This yields an mfwmax/gl - ratio of 0. 1 .

The anterior extremity of thè gladius is sharply pointed.

From posterior to anterior a constantly 0.7 mm thick

rounded massive keel is prominent on thè median field.

Imprints of thè ventral gladius surface in thè anterior part

(where gladius material is missing) show that thè median

keel is ventrally two-parted (see cross-section Fig. 5; PI.

IV, Fig. B). Since these structures are not present on thè

dorsal surface, thè ventral surface differs from thè dorsal

surface. On both sides of thè keel lateral ridges form out-

ermost margins. These are also counter drawn on thè ven¬

tral surface, as shown by imprints where gladius material

is not preserved (PI. IV, Fig. B).

The lateral fields appear very thin, marginally even

translucent. Growth lines on thè dorsal surface of both

lateral fields can be easily traced but exhibit just a weakly

curved hyperbolar zone (PI. IV, Fig. C). This implies that

outer asymptotes are present but inconspicuous. Inner

asymptotes divide thè lateral and thè median field and are

thus well marked by lateral ridges. Lateral ridges diverge

at an angle of 8°.

Arms: as already mentioned, especially Paratype I

(Fig. 6; PI. V, Fig. A; PI. VI, Fig. B) displays an arm

crown contrasting with thè surrounding colour of sedi-

ment (surprisingly they are not preserved as phosphatised

musculature). At least six arms can be counted. A seventh

arm is only visible under ultraviolet light (PI. VI, Fig. B).

They are all uniform and slender (approximately 5 cm in

length and 0.3 mm in diameter).

In Paratype II (PI. V, Fig. B; PI. VI, Fig. C), arms

are preserved as 3-dimensional ridges similar to those

described from Miinstere/la scutellaris (Fuchs et al.,

2003: 4). As in lateral specimens of M. scutellaris, ultra¬

violet light reveals thè phosphatised musculature of four

arms, which are considered to be thè right arms of each

pair. There is no indication of an arm web between thè

Fig. 6 - Rachiteuthis donovani. Drawing of Paratype I (MSNM i25 135).

ac = arm crown; bm = buccal mass; f = fin; fu = funnel; g = gladius; is

- ine sac.

10

DIRK FUCHS

ridges. At least thè inner surface of one left arm is pre-

served (most probably thè ventrolateral arm). Tentacles,

tentacle clubs, suckers or cirri are not observable in any

of thè specimens.

Head: neither in thè Holotype nor in thè Paratypes is

there any reason to suppose a clearly demarcated head. In

both Paratypes it seems that dorsal arm bases are a direct

continuation of thè dorsal mantle edge (Pls. V and VI).

A head-mantle fusion would imply that thè head was not

clearly demarcated.

From specimens of Miinsterella scutellaris (Fuchs et

al, 2003: 4) it was demonstrated that head retraction was

possible in this taxon. Similar abilities are assumed for

Rachiteuthis. The arm bases of both Paratypes He very

dose to thè anterior end of thè gladius, which should

mark thè anterior mantle edge. The buccal mass of Para-

type I is obviously situated within thè mantle (PI. V, Fig.

A). Subsequently thè head must have been retracted.

This is not thè case in thè Holotype, where thè head is

apparently a short distance in front of thè anterior end of

thè gladius and thus presumably stretcbed out. A head

retractor muscle, which is observed in Miinsterella, is not

preserved.

Beaks: a weak parabolic structure within thè head

region of thè Holotype is determined as a lower or upper

beak but contains no reai morphological information as

obtained from Trachyteuthis, Plesioteuthis or Lepto-

teuthis from Nusplingen (Klug et al., 2005).

Body outline: exceptional preservation of Paratype

I allows statements about thè complete body outline

of Rachiteuthis (PI. V, Fig. A). The body was appar¬

ently twisted during diagenesis. Although thè funnel

and soft-body are laterally embedded, gladius and fins

are seen in dorsal view. During decay, thè stiff gladius

must have penetrated thè anterior mantle. However, it

is assumed that thè body was nearly circular in cross-

section and comparatively longish and slender (tor-

pedo-shaped).

Funnel: funnel preservation is extremely rare even in

Konservat-Lagerstàtten. In Paratype I (Fig. 6; PI. V, Fig.

A) a structure determined as thè fiinnel is situated beneath

thè anterior part of thè gladius. It is 0.6 cm in diameter.

The length is difficult to define. It is obvious that ink was

ejected through thè funnel. Like thè head, thè funnel

seems to be retracted.

Fins: although phosphatised musculature is missing in

Paratype I, posteriorly a triangular outline of paired fins

is clearly visible (Fig. 6; PI. V, Fig. A). The outer corners

are distinctly rounded. They begin with thè posteriormost

extremity of thè gladius and end 4.5 cm anterior to this

point. Their rounded margins are 4.8 cm apart.

Differential diagnosis: gladius characteristics such

as opened conus with comparatively short lateral fìelds,

presence of a hyperbolar zone on thè lateral fìelds as well

as a distinctly pointed anterior end justify thè taxonomic

position within thè Teudopsidina.

The presence of a median keel and paired lateral

ridges in connection with a shaft-like prolongation of thè

rachis-like median field resembles Palaeololigo oblon-

gata Wagner, 1 859 or P. albersdoerferi Engeser & Keupp,

1999 from thè Lithographic Limestones of Solnhofen. In

contrast, thè free rachis of Rachiteutis is much longer and

thè hyperbolar zone is less distinct than in Palaeololigo.

A median keel and lateral ridges also occur in forms like

Plesioteuthis and Dorateuthis. Compared to these forms

lateral fìelds of Rachiteuthis do not form a closed conus

but are clearly spread out.

Family Trachyteuthididae Naef, 1921

Genus Trachyteuthis v. Meyer, 1 846

(= Coccoteuthis Owen, 1855; Voltzia Schevill, 1950;

junior subjective synonyms)

Type species: Sepia hastiformis Riippell, 1829 (by

subsequent designation (Bulow-Trummer, 1920: 248).

Geographic and stratigraphic distribution: previous

records of thè genus Trachyteuthis are from thè Callovian

of Christian Malford, England (Martill & Hudson, 1994),

thè Oxfordian of Chile (Schulze, 1989), Graham Land

of Antarctica (Doyle, 1991) and Vinales region of Cuba

(Schevill, 1950), thè Upper Kimmeridge of Dorset, Eng¬

land (Donovan, 1977) and Nusplingen, Germany (Klug

et al., 2005), thè Tithonian of Solnhofen and thè Volga

region in Russia (Hecker & Hecker, 1955), thè Aptian of

Northwest Queensland, Australia (Wade, 1993) and Heli-

goland, Germany (Engeser & Reitner, 1985).

As thè following descriptions show, we are now able

to extend thè stratigraphical range of Trachyteuthis at

least up to thè Upper Cenomanian.

Remarks: many workers have considered this genus to

be a decabrachian, because of granulation on thè dorsal

surface of thè gladius similar to recent cuttlebones of

Sepia (Donovan, 1977; Doyle, 1991). Despite hundreds

of well-preserved specimens from Nusplingen and Sol¬

nhofen there are no indications for a chambered complex

associated with thè gladius. Additionaly, specimens with

ten arms have never been observed. Finally, Donovan

(2002) presented a specimen of Trachyteuthis with two

pairs of fins. This character is only shared with thè recent

Vampyroteuthis inferna/is. Except granulation, charac-

teristic features for a decabrachian affiliation are absent.

Therefore Trachyteuthis must ultimately be treated as a

reai representative of thè Vampyropoda (Bandel & Leich,

1986; Doyle et al., 1994; Donovan, 2002; Haas, 2002;

Bizikov, 2004).

Trachyteuthis cf. hastiformis Riippel, 1829 (PI. VII)

Holotype: Senckenberg Museum, Frankfurt/Main (Ge¬

rmany), register number XI 1328.

Locus typicus: Lithographic Limestones of Solnho¬

fen.

Stratum typicum: Lithographic Limestones of Solnho¬

fen, Lower Tithonian (Malm, £ 2b).

Geographic and stratigraphic distribution: as for thè

genus (?).

Material: MSNM Ì20591 (PI. VII), MSNM Ì20590 and

MSNM Ì25145 come from thè locality of Hàqel.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

11

Description

In each specimen parts of thè gladius as well as mantle

musculature are preserved. Neither ink sac, fin, head nor

arm crown are observable in any of thè specimens.

In MSNM Ì25145 mainly muscular mantle is pre¬

served. The total length of this specimen is 26 cm. In most

places musculature traces thè outline of thè anterior part

of thè gladius. Marginai remains of thè gladius are stili

present at thè anteriormost end, and one can follow thè

characteristically pointed anterior margin of thè Trachy-

teuthis gladius. At a distance of 14 cm from this margin at

a small spot in thè median line, characteristic growth lines

are “shining” through thè musculature. Specimen MSNM

Ì20590 is in such a poor condition that its identification as

Trachyteuthis is merely presumed.

Specimen MSNM Ì20591 (Fig. 7; PI. XII) displays a

complete outline of thè gladius. The gladius was embed-

ded dorsally and is seen in ventral view. The majority of

gladius material is originally preserved. Imprints of thè

dorsal surface are visible, where gladius material is miss-

ing. Measurements are as follows:

- gladius length (gl): 36.0 cm

- lateral field length (lfl): 14.5 cm

- median field width (mfwif) at thè anterior end of thè

lateral fields: 9.0 cm

- maximum lateral field width (lfwmax): 13.0 cm

- lateral field length/gladius length (lfl/gl): 0.4

- maximum median field width/gladius length (mfwmax/

gl): 0.25

- maximum lateral field width/lateral field length (lfwmax/

lfl): 0.9

Owing to thè ventral view there is nothing visible from

thè typical dorsal granulation. However, at thè posterior-

most end of thè gladius weak imprints of granulations are

visible similar to those of Trachyteuthis hastiformis from

Solnhofen and Nusplingen.

Although thè ends of thè lateral fields are well pre¬

served, a hyperbolar zone (as thè transition from thè

lateral fields into thè median field) is indistinct. As a

result inner and outer asymptotes are merely conjec-

turable. Inner asymptotes diverge at an apical angle of

approximately 37°. Inner asymptotes are commonly

defined as thè end of thè lateral fields. Diverging struc-

tures on thè median field should not be mistaken for

inner asymptotes. The area between these structures is

considered to be thè granulated part of thè median field.

This granulated median part is stronger developed than

marginai parts of thè median field and diverges at an

angle of approximately 10°. A well-preserved dorsal

view of a Trachyteuthis- gladius from Nusplingen, illus-

trated in Klug et al. (2005), displays an identical dorsal

strengthening.

Each ratio enumerated above as well as thè apical

angle fits well within thè ranges known from specimens

from Solnhofen and Nusplingen.

Differential diagnosis: present specimens are unam-

biguous representatives of thè genus Trachyteuthis and

are very dose to T. hastiformis from Solnhofen and

Nusplingen. Compared to a dose relative from Hàqel and

Sàhel Aalma, Glyphiteuthis ( =Libanoteuthis , see below),

Trachyteuthis cf. hastiformis differs in its much bigger

dimensions and thè lack of a median keel.

dorsal view posterolateral view

Fig. 7 - Trachyteuthis cf. hastiformis. Reconstruction of thè gladius. cl = conus length; ga = granulated area; gl = gladius length; hz =

hyperbolar zone; ia = inner asymptote; lf = lateral field; lfl = lateral field length; lfw = lateral field width; mf = median field; mfw =

median field width; oa - outer asymptote.

12

DIRK FUCHS

Remarks: an unusual high number of specific names

exist in thè literature for forms of Trachyteuthis hasti-

formis from thè Solnhofener Plattenkalks. Although

Engeser (1988) regarded them altogether in synonymy,

thè presence of more than only one morphospecies cannot

be ruled out (Fuchs, 2006). Further, species of thè genus

Trachyteuthis from other locations than Solnhofen cannot

be clearly distinguished from T. hastiformis by means of

adequate characters. Finally, there are no distinct differ-

ences in proportions even between Jurassic and Creta-

ceous examples. Since thè fìrst occurrence of thè genus

in thè middle Jurassic no modifications are observable at

fìrst glance. As already mentioned by Donovan (1995) a

comparative examination is necessary in order to specify

morphological differences. Therefore, these specimens are

provisionally determined as Trachyteuthis cf. hastiformis.

Genus Glyphiteuthis Reuss, 1854 (Pls. VIII-IX)

Type species: Glyphiteuthis ornata Reuss, 1854.

Geographic and stratigraphic distribution: Lower/

Middle Turonian of Bilà Hora Formation (Czech Repub-

lic), Upper Cenomanian of Lebanon and Normandy

(France), Upper Santonian of Sàhel Aalma (Lebanon).

Remarks: previously nobody has referred to thè dif-

ficult morphological distinction between thè Turonian

genus Glyphiteuthis Reuss, 1854 from Bohemia (Czech

Republic) and Libanoteuthis Kretzoi, 1942 from Hàqel,

Hjoula and Sàhel Aalma. Kretzoi (1942) erected Liba¬

noteuthis without any descriptive explanations. While

Roger (1946), Rioult (1962), Waage (1965), Engeser &

Reitner (1986) and Engeser (1988) have used this genus

name, Jeletzky (1966) and Donovan (1977) have placed

this taxon within thè genus Trachyteuthis. Certainly, there

are enough morphological differences between thè genera

Libanoteuthis and Trachyteuthis. Especially thè anterior

end of thè Trachyteuthis-gìadius is completely different

from Libanoteuthis. Instead, dose affinities to Glyphi¬

teuthis should be discussed. Re-investigations of speci¬

mens of Glyphiteuthis ornata (specimens of G. minor are

considered to be juveniles of G. ornata) and specimens of

“ Libanoteuthis ” libanotica yielded equal gladius propor¬

tions. In thè Turonian specimens it seems that thè median

ridge is more developed than in Cenomanian specimens.

However, this is possibly due to thè flattened specimens

of thè Lebanese “fish beds” in comparison to a 3-dimen-

sional preservation in thè Bilà Hora Formation (Bohemia).

In any case, there are only few arguments for separating

specimens of Glyphiteuthis and Libanoteuthis in different

genera. Therefore Libanoteuthis should be considered as

a junior synonym of Glyphiteuthis.

Assigned species: Glyphiteuthis ornata (= minori),

G. libanotica and G. boutillieri.

Glyphiteuthis (= Libanoteuthis) libanotica Fraas, 1878

(PI. Vili, Figs. A-B; PI. IX)

1878 Geoteuthis libanotica - Fraas, p. 345, PI. 6, Fig. 3.

1896 Geoteuthis libanoticus - Woodward, p. 23 1 .

1920 Geoteuthis libanotica Fraas - Bulow-Trummer, p.

255.

1922 Trachyteuthis libanotica (Fraas, 1878)-Naef, p. 140,

Fig. 49b.

1942 Libanoteuthis libanotica Fraas - Kretzoi, p. 125,

Fig. 1.12, p. 134.

? 1 946 Geoteuthis sahel-alma (Fraas) (Naef) - Roger, p. 1 6,

Fig. 9.

1946 Trachyteuthis libanotica (Fraas) - Roger, p. 17, Fig.

10.

1952 Libanoteuthis libanotica Fraas - Roger, p. 740.

1 962 Libanoteuthis libanotica (Fraas) - Rioult, p. 1 1 , Fig.

lb.

1966 Trachyteuthis (Libanoteuthis) libanotica Kretzoi,

1942 - Jeletzky, p. 45.

1977 Trachyteuthis libanotica (Fraas) - Donovan, p. 32.

1986 Libanoteuthis libanotica (Fraas, 1878) - Engeser &

Reitner, p. 7, Fig. 2, PI. 2, Figs. 1, 2.

1988 Libanoteuthis libanotica (Fraas, 1878) - Engeser,

p. 69.

Holotype: Fraas (1878: PI. Vili, Fig. B), Staatliches

Museum fur Naturkunde, Stuttgart (SMNS 26271).

Locus typicus: Hàqel, Lebanon.

Stratum typicum: Upper Cenomanian.

Geographic and stratigraphic distribution: Upper Ce¬

nomanian of Hàqel and Hjoùla, Upper Santonian of Sàhel

Aalma, (Lebanon).

For a long time Glyphiteuthis libanotica has been known

only from Hàqel, but Engeser & Reitner (1986: 9) consid¬

ered thè possibility that Naef (1922) mistook thè localities

of his specimens from thè Noetling collection (Noetling,

1 886), which were believed to derive from Sàhel Aalma.

This implies that this genus occurs up to thè Santonian.

Material: eighteen specimens identified as Glyphi¬

teuthis libanotica belong to thè MSNM-collection and

were investigated: MSNM i25 138 (PI. IX, Fig. A), Ì24801

(PI. IX, Fig. B), Ì25127 (PI. Vili, Fig. A), Ì24959, Ì20589,

Ì24957, Ì20588, i20857a+b, Ì25136, Ì25130, Ì12599,

Ì25137, i25131a+b, Ì24958, Ì23434, Ì25129, Ì26734,

Ì23422. Except MSNM Ì2480, which comes from Hjoùla,

all specimens originate from Hàqel.

Description

Gladius: dimensions of eight dorsoventrally embed-

ded specimens are completely preserved and are therefore

available for comparative measurements (Fig. 8).

With gladius lengths of 4-8 cm, all measured speci¬

mens are small compared to Glyphiteuthis ornata and

Trachyteuthis. Lateral field length (lfl) is between 1.4-3. 4

cm. The ratio of lateral fìeld length to gladius length (lfl/

gl) varies slightly between 0.36-0.4. Maximum width of

thè lateral field (lfwmax) is between 1.3-2. 6 cm. In cross-

section thè lateral fields are almost hemi-circular. The

ratio maximum lateral field width to lateral field length

is between 0.9-1. Especially specimen MSNM i25 1 27

exhibits a precise gladius outline (PI. Vili, Fig. A). This

specimen clearly shows a sharply pointed anterior end as

well as a slight lateral constriction in thè anterior third of

thè gladius.

In specimens MSNM i25 1 30, Ì24958 and Ì24959 an

indistinct but prominent median ridge or even keel (“Mittel-

linie” of Engeser & Reitner (1986: 7) is present. In contrast

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

13

dorsal view

Fig. 8 - Glyphiteuthis (= Libanoteuthis ) libanotica. Reconstruction

of thè gladius. ga= granulateci area; hz = hyperbolar zone; ia = inner

asymptote; lf = lateral field; lfl = lateral field length; mf = median field;

mk = median keel; oa = outer asymptote.

to observations by Engeser & Reitner (1986) thè median

ridge runs continuously from posterior to anterior. After re-

investigation of thè Holotype of Glyphiteuthis libanotica it

must be assumed that thè lack of a posterior median ridge

in this specimen is due to incomplete preservation.

MSNM Ì24959 displays dorsal imprints of granula-

tions. In this specimen one row of coarse granules is

visible along thè median ridge. Smoother granules flank

thè median ridge on both sides. Additional ridges similar

to those assumed in Trachyteuthis cf. hastiformis (see

above) strictly delimit thè granulation. They diverge

with an angle of 20°. As in Trachyteuthis cf. hastiformis

it seems that thè granulated area in MSNM i25 138 is

slightly elevated above thè rest of thè gladius surface (PI.

IX, Fig. A). In thè middle part of thè gladius thè ridges

become indistinct and thè granulation broadens. In thè

anterior third of thè gladius it almost reaches thè marginai

edges (MSNM Ì20589).

Preservation of growth lines is weak (MSNM i25 138).

In some specimens imprints of growth lines are preserved

(MSNM Ì24958). It seems that growth lines have a weak

curvature, and so thè hyperbolar zone is indistinct. Inner

and outer asymptotes are therefore imaginary.

Arms: in at least ten specimens (MSNM Ì24957,

Ì24958, Ì25129, i25131a+b, Ì24959, Ì24801, Ì25130,

Ì25138, i20587a+b, il 2599) arms are more or less pre¬

served as outstanding discolorations. Phosphatised arm

musculature is only preserved in MSNM Ì20875 but

counting has proved to be difficult. Although a complete

arm crown is absent, each specimen suggests that these

animals had relatively long and slender arms (PI. IX, Fig.

B). Average arm thickness is 0.2 mm. Absolute arm length

could not be determined. Tentacles or tentacle clubs are

never present.

Suckers: few imprints of suckers are discemible in

specimen MSNM i25 130. It is not possible to determine

thè position of suckers (for instance, if they are arranged

in one, two or more rows). Presumably they form distai

suckers. They are almost 1 mm in diameter and radiai in

shape. Cirri are never observed.

Head: in each specimen where remains of arms are

visible remains of thè head can be assumed by thè posi¬

tion of thè buccal mass.

At least two specimens (MSNM il 2599, in ventral

view and Ì24801, in lateral view) exhibit a circular whit-

ish structure in thè head region, strongly phosphatising

under ultraviolet light (PI. IX, Figs. B-C). This structure is

determined as thè muscular buccal mass. In both cases it

almost touches thè anterior end of thè gladius. This might

indicate a retractable head. Other specimens confimi this

assumption. However, it can not be decided if thè head

was clearly demarcated.

Beaks: beaks occur in several specimens but provide

no information about their detailed morphology owing to

strong compaction.

Fins: absolutely no indications about fin morphology

are present, neither in specimens of this collection nor in

specimens from other collections.

Remarks: as mentioned above thè ratio lateral field length

to gladius length (lfl/gl) varies between 0.36-0.40. The

Holotype of Glyphiteuthis libanotica was measured by thè

author and thè lfl/gl-ratio of 0.38 from thè Holotype fits well

into this range. It must be emphasised that measurements

of flattened (originally 3-dimensional) structures always

contain errors because thè rates of flattening usually differ.

Nevertheless, at least one dorso ventral ly flattened speci¬

men (MSNM Ì25131) with a lateral field length to gladius

length ratio (lfl/gl) of 0.3 gives thè impression that probably

two morphotypes occur. Though this cannot be statistically

proved, it seems that there is 1) a stouter form with circa

38% and 2) a more slender form with lateral field length

of circa 30% of thè gladius length. Possibly arm length is a

second distinguishing character, but present specimens do

not provide sufificient infonnation. Additional gladii with

well-preserved soft-parts are therefore necessary.

Differential diagnosis: according to Engeser & Reitner

(1986) co-existing G. boutillieri differ from G. libanotica

by a solid median ridge and a specific median field. As

shown above in G. libanotica thè median ridge is con-

tinuous, too. Judging from thè drawings of Lennier (1886)

and Rioult (1962) likewise no difference conceming thè

median field is recognizable. Riegraf (1987) takes into

consideration that G. libanotica is a junior synonym of G.

boutillieri. In order to define specific differences between

these two forms a re-investigation of thè Holotype of G.

boutillieri is necessary.

G. ornata possesses a ribbed median keel especially

in thè anterior third of thè gladius (Kostak, 2002). In G.

libanotica we have found no indications for ribs.

14

DIRK FUCHS

PHYLOGENETIC IMPLICATIONS

Since evidences for a dose affiliation of fossil gladii

with recent Vampyroteuthis and octopods have accumu-

lated in recent decades (Bandel & Leich, 1986; Haas,

2002; Donovan et al., 2003; Bizikov, 2004; Fuchs, 2006),

“fossil teuthids” are unknown in thè fossil record.

According to Engeser & Reitner (1986), Bandel &

Leich (1986) and Engeser (1988) Prototeuthina, Loli-

gosepiina and Teudopsidina (= Mesoteuthina) represent

stem-lineage representatives of thè Vampyromorpha.

In contrast Haas (2002) considers them as stem-line-

age representatives of all Vampyropoda. Bizikov (2004)

assumes that thè Loligosepiina are thè stem group of thè

Vampyropoda and that teudopsid forms are stem-lineage

representatives of thè Octobrachia.

However, on thè basis of hundreds of vampyropod

gladii from different localities and stratigraphic levels a

comparative study suggests that only thè Prototeuthina

(Paraplesioteuthis, Plesioteuthis, Dorateuthis) should be

regarded as stem-lineage representatives of thè Vampyro¬

poda (Fuchs, 2006). Most probably thè ventrally opened

conus of thè Loligosepiina and thè Teudopsidina is a

derivation of a closed conus (as in Paraplesioteuthis and

Plesioteuthis) inherited from a phragmoteuthid ancestor

(Fig. 9).

Compared to thè Loligosepiina, Teudopsidina show

a generai tendency to reduce their median fields (Haas,

2002; Bizikov, 2004). Since thè gladius rudiments of

incirrate octopods ( Palaeoctopus newboldi) are proved

to be thè result of a progressive reduction of thè median

field, Teudopsidina should indeed be considered as stem-

lineage representatives of thè Octobrachia.

Recent Vampyroteuthis infernalis exhibits gladius

characteristics that can easily be derived from a loli-

gosepiid gladius, which is why Loligosepiina are believed

to be thè stem group of thè Vampyromorpha (Fuchs,

2006).

It must be pointed out that thè Lithographic Lime-

stones of Hàqel, Hjoula and Sàhel Aalma apparently do

not contain unambiguous representatives of thè Decabra-

chia despite its extraordinary preservational potential.

VAMPYROMORPHA CIRROCTOPODA OCTOPODA

Fig. 9 - Basic phylogeny of thè Vampyropoda.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

15

CONCLUSIONS

Although well-preserved coleoids from Lebanon

were already praised during thè nineteenth century, pre-

vious information about diversity and morphology was

scarce. The great number of Lebanese coleoids enables

to enlarge thè diversity up to six species ( Acanthoteuthis

syriaca, Palaeoctopus newboldi, Dorateuthis syriaca,

Rachiteuthis donovani, Trachyteuthis cf. hastiformis,

Glyphiteuthis libanotica ) and to increase thè morphologi-

cal understanding of most of them.

Hàqel, Hjoula, and Sàhel Aalma exhibit a vampyropod

diversity that consists of a mixture of conservative and

modified species. Especially thè discovery of Trachy¬

teuthis demonstrates a remarkable evolutionary stability

within this genus from thè Jurassic up to thè Late Creta-

ceous. Glyphiteuthis displays only a slight modification of

a typical teudopsid gladius.

The gladius of Rachiteuthis can be seen as a modifìca-

tion of a palaeololiginid gladius. Whereas thè posterior

part of thè gladius with its lateral fields is more or less

unmodified thè rachis-like median fìeld is strongly elon-

gated compared to Palaeololigo.

The gladius of Dorateuthis must be considered as

a moderate modification, which derived from a Ple-

sioteuthis-Mke gladius. Plesioteuthis possesses a reduced

but ventrally closed conus with short lateral fields. The

median field is comparatively narrow but well developed.

In Dorateuthis, in contrast, thè median field seems to

be only membranous with a very poor preservational

potential. In this way thè sturdy lateral ridges exclusively

provide for thè construction of thè gladius and thus for thè

longitudinal rigidity of thè mantle cavity. Morphological

knowledge about thè conus of Dorateuthis is poor, but it

seems that it is distinctly reduced.

Except in extraordinary Konservat-Lagerstàtten such

as Christian Malford, Holzmaden, Solnhofen and Leba¬

non, coleoid records are often sporadic (except belemnite

rostra). Owing to bad preservational conditions, morpho¬

logical comparisons between fragmentary specimens are

usually problematic. This remarkable coleoid collection

from Hàqel stored in thè Museo di Civico di Storia Natu¬

rale di Milano demonstrates thè advantages and espe¬

cially thè importance of a suffìcient number of specimens

in order to verify earlier assumptions and to present a reli-

able morphological and taxonomical evaluation.

Unfortunately, an adequate Konservat-Lagerstàtte is

missing between thè “evolutive Windows” Solnhofen

(Tithonian) and Sàhel Aalma (Santonian). Sàhel Aalma

forms thè last “evolutive window” in thè earth history

for we do not have any Konservat-Lagerstàtte containing

coleoids during thè Cenozoic (coleoids are apparently

unknown in Monte Bolca). Present results demonstrate

thè enormous significale of Hàqel, Hjoùla, and Sàhel

Aalma for thè reconstruction of thè evolutionary history

of vampyropods. Interestingly, no information about cole¬

oids from Nàmmoura is available.

Most probably, present contributions are preliminary,

because additional information about Lebanese coleoids

is expected from another 80 specimens housed in thè

Black Hills Museum of Naturai History in Hill City

(South Dakota, USA).

Acknowledgments

Present work was funded by thè Deutsche Forschungs-

gesellschaft (DFG). Special thanks go to A. Garassino and

G. Teruzzi (both from thè Museo Civico di Storia Naturale

di Milano) who enabled me to investigate thè collection.

I am grateful to Neal L. Larson (Black Hills Institute)

for his thorough reading of thè manuscript as well as his

constructive suggestions for its improvement.

16

DIRK FUCHS

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Dirk. 1 uchs - Freie Universitàt Berlin, Institut tur Geologische Wissenschaften, Fachrichtung Palàontologie, Malteserstr. 74-100,

D- 12249 Berlin, Germany.

e-mail: drig@zedat.fu-berlin.de

Morphology, taxonomy and diversity of vampyropod Coleoids (Cephalopoda) from thè Upper Cretaceous of Lebanon

Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano

Volume XXXI V - Fascicolo II

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

19

mm

PI. I - Dorateuthis syriaca. A) Specimen MSNM i25 1 44 from Hjoùla. Seen in ventral view. is = ink sac; Ir = lateral ridge; se = stomach

contents. Scale bar: 1 cm. B) Same specimen as in Fig. A. Close up of thè posterior region. Punctuated line marks supposed lateral

fields (10; Ir = lateral ridge. Scale bar: 1 cm. C) Specimen MSNM i23 1 08 from Sàhel Aalma. Seen in ventral view. Ir = lateral ridge;

mf = median field; mm = mantle muscolature; mr = median ridge. Scale bar: 1 cm.

20

DIRK FUCHS

PI. II - Dorateuthis syriaca. A) Specimen MSNM Ì24800 from Hjoùla. Seen in lateral view. bm = buccal mass; co = conus. Arrow

heads mark and delimit thè outline of thè ventral mantle. Scale bar: 1 cm. B) Specimen MSNM i25 1 33 from Hjoùla. Seen in lateral

view. ac = arm crown; bm = buccal mass; is = ink sac. Scale bar: 1 cm. C) A specimen from Sàhel Aalma deposited in thè Naturhis-

torisches Museum Wien, NHMW 1998z0 105/0000 (originai of Lukeneder & Harzhauser 2004). Seen in ventral view. The arm crown

consists ot eight arms. Note, that thickness of arms, body as well as gladius outline is a preparational artefact. Scale bar: 1 cm.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

21

( <4v^y/

' ■ ^ Ì8L

PI. Ili - Dorateuthis syriaca. A) Specimen MSNM i25 1 34 from Hjoùla. Seen in lateral view. b = beaks; bm = buccal mass; f = fin; is = ink sac.

Scale bar: 1 cm. B) Holotype of Dorateuthis syriaca from Sàhel Aalma (British Museum of Naturai History London, BMNH C501 7). On thè

left a weak arm crown is visible. On thè right weak imprints of thè gladius with lateral and median ridges. Scale bar: 1 cm. C) Holotype of

Dorateuthis sahilalmae from Sàhel Aalma (Staatliches Museum fìir Naturkunde Stuttgart, SMNS 26269). Seen in dorsal view. Lateral ridges

(Ir) diverge at a remarkable apikal angle. Most probably this due to a longitudinal disrupted gladius. is = ink sac. Scale bar: 1 cm.

22

DIRK. FUCHS

PI. IV - Rachiteuthis donovani ( nov ; spec.). A) Holotype (MSNM i25 1 42) from Hjoùla. Seen in dorsal view. Scale bar: 1 cm. B) Sartie

specimen as in Fig. A. Close up ot thè anterior part of thè gladius. Where parts of thè gladius have not been preserved, imprints of thè

ventral side are visible. id = ink duct; ntk = ntedian keel; mm = muscular mantle. Scale bar: 1 cm. C) Same specimen as in Fig. A. Close

up ot thè posterior part ot thè gladius. hz = hyperbolar zone; lf = lateral field; Ir = lateral ridge; mk = median keel. Scale bar: 1 cm.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

23

’&sa

bm

ac

r .. JjK *

tf, ' r.

v4jM-

_

PI. V - Rachiteuthis donovani ( nov : spec.). A) Paratype I (MSNM Ì25135) from Hjoula. The gladius is seen in dorsal view, whereas thè

body is in lateral aspect. ac = arm crown; bm = buccal mass; f = fin; fu = funnel; is = ink sac. Scale bar: 1 cm. B) Paratype li (MSNM

Ì25139) from Hjoula. Gladius in dorsal view. Body in lateral view. is = ink sac. Scale bar: 1 cm. Left bottoni corner: a dose up of thè

head region. It seems as if thè dorsal mantle is fused with thè head region.

24

DIRK FUCHS

M- \ 1 ■ Rachìteuthis donovani ( nov : spec.) photographed with ultra violet light. A) Holotype. Phosphatised remains are shining white.

ruma posterior mantle margin. Scale bar: 1 cm. B) Paratype I. Although tissues are poorly posphatised, number of arms (1-8), body

as well as fin outlines are distinctive. Scale bar: 1 cm. C) Paratype II. At least four arm stubs are visible. Scale bar: 1 cm.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

25

PI. VII - Trachyteuthis cf. hastiformis. A) Specimen MSNM Ì20591 from Hàqel. Seen in ventral view. Gladius outline is clearly visible.

Soft parts are not preserved. Scale bar: 1 cm. Fig. B) Same specimen as in Fig. A. Close up of thè anterior part. ga = granulated area.

Scale bar: 1 cm. Fig. C) Same specimen as in Fig. A. Close up of thè posterior part. ga = granulated area; hz = hyperbolar zone; ia =

inner asymptote; lf = lateral field; mf = median field; oa = outer asymptote. Scale bar: 1 cm.

26

DIRK FUCHS

PI. VII! - Glyphiteuthis. A) Glyphiteuthis ( Libcinoteuthis ) libanotica from Hàqel. Specimen MSNM i25 127. Seen in dorsal view. Scale

bar: 1 cm. B) Holotype of Glyphiteuthis ( Libanoteuthis ) libanotica (Staatliches Museum fìir Naturkunde Stuttgart, SMNS 26269)

from Hàqel. Seen in dorsal view. Scale bar: 1 cm. C) Holotype of Glyphiteuthis ornata (National Museum Prague, 06099) from thè

Turonian of Bohemia (CZ). Seen in dorsal view. Scale bar: 1 cm.

MORPHOLOGY, TAXONOMY AND DIVERSITY OF VAMPYROPOD COLEOIDS (CEPHALOPODA) FROM THE UPPER CRETACEOUS OF LEBANON

27

PI. IX - Glyphiteuthis libanotica. A) Specimen MSNM i25 1 38 from Hàqel. ga = granulateci area; hz = hyperbolar zone; is = ink sac; ia

= inner asymptote; lf = lateral field; mf = median field; oa = outer asymptote. Seen in dorsal view. Scale bar: 1 cm. B) Specimen MSNM

Ì24801 from Hàqel. Seen in lateral view. ab = armbases; bm = buccal mass; mm = mantle muscolature. Scale bar: 1 cm. C) Same speci¬

men as in Fig. B photographed with ultra violet light. ab = armbases; bm = buccal mass; mm = mantle musculature. Scale bar: 1 cm.

28

DIRK FUCHS

INDEX

Introduction . Pag. 3

Previous studies on coleoid cephalopods from

thè Late Cretaceous of Lebanon . Pag. 4

Geological setting . Pag. 4

Material . Pag. 5

Modes of preservation . Pag. 5

Systematic palaeontology . Pag. 6

Phylogenetic implications . Pag. 14

Conclusions . Pag. 15

Acknowledgments . Pag. 15

References . Pag. 16

l

Ili - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Al¬

pe Turati (Erba, Como). Generi Hildoceras, Phymatoceras, Paronice-

ras e Frechiella. Conclusioni generali./?/?. 143-204, 2 figg., 6 tavv.

Volume XVIII

I - PINNA G., 1969 - Revisione delle ammoniti figurate da Giuseppe Me¬

neghini nelle Tavv. 1-22 della «Monographie des fossiles du calcaire

rouge ammonitique » ( 1 867- 1881)./?/?. 5-22, 2 figg. , 6 tavv.

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

d’Orta )./?/?. 23-92, 42 figg., 4 tavv. n.t.

Ili - PETRUCCI F., BORTOLAMI G. C. & DAL P1AZ G. V., 1970 - Ri¬

cerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul

suo substrato cristallino. /?/?. 93-169, con carta a colori al 1:40.000, 14

figg., 4 tavv. a colori e 2 b.n.

Volume XIX

I - CANTALUPPI G., 1 970 - Le Hildoceratidae del Lias medio delle regio¬

ni mediterranee - Loro successione e modificazioni nel tempo. Riflessi

biostratigrafici e sistematici./?/?. 5-46, 2 tabb. n.t.

II - PINNA G. & LEVI-SETT1 F„ 1971 - 1 Dactylioceratidae della Provin¬

cia Mediterranea (Cephalopoda Ammonoidea). pp. 47-136, 21 figg.,

12 tavv.

Ili - PELOSIO G., 1973 - Le ammoniti del Trias medio di Asklepieion (Ar-

golide, Grecia) - 1. Fauna del «calcare a Ptychites » (Anisico sup.). pp.

137-168, 3 figg., 9 taw.

Volume XX

I - CORNAGGIA CASTIGLIONI 0., 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./?/?. 81-127,

37 figg., 6 carte tematiche.

Ili - CERETTI E. & POLUZZI A., 1973 - Briozoi della biocalcarenite 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 figg., 16 tavv.

II - POLUZZI A., 1975 - 1 Briozoi Cheilostomi del Pliocene della Val d’Ar-

da (Piacenza, Italia)./?/?. 35-78, 6 figg., 5 tavv.

Ili - BRAMBILLA G., 1976 - I Molluschi pliocenici di Villalvernia (Ales¬

sandria). I. Lamellibranchi. pp. 79-128, 4 figg., 10 taw.

Volume XXII

I - CORNAGGIA CASTIGLIONI O. & CALEGARI G„ 1978 - Corpus

delle pintaderas preistoriche italiane. Problematica, schede, iconogra¬

fia./?/?. 5-30, 6 figg., 13 tavv.

II - PINNA G., 1979 - Osteologia dello scheletro di Kritosaurus notabilis

(Lambe, 1914) del Museo Civico di Storia Naturale di Milano (Ornithi-

schia Hadrosauridae). pp. 31-56, 3 figg., 9 tavv.

Ili - BIANCOTTI A., 1981 - Geomorfologia dell’Alta Langa (Piemonte

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

Volume XXIII

I - GIACOBINI G., CALEGARI G. & PINNA G., 1 982 - 1 resti umani fos¬

sili della zona di Arena Po (Pavia). Descrizione e problematica di una

serie di reperti di probabile età paleolitica./?/?. 5-44, 4 figg., 16 tavv.

II - POLUZZI A., 1982 - 1 Radiolari quaternari di un ambiente idrotermale

del Mar Tirreno, pp. 45-72, 3 figg., 1 tab., 13 taw.

Ili - ROSSI F., 1984 - Ammoniti del Kimmeridgiano superiore-Berriasiano

inferiore del Passo del Furio (Appennino Umbro-Marchigiano), pp. 73-

138, 9 figg., 2 tabb., 8 taw.

Volume XXIV

I - PINNA G., 1 984 - Osteologia di Drepanosaurus unguicaudatus, lepido-

sauro triassico del sottordine Lacertilia. pp. 5-28, 12 figg., 2 taw.

II - NOSOTTI S. e PINNA G., 1989 - Storia delle ricerche e degli studi

sui rettili Placodonti. Parte prima 1830-1902. pp. 29-86, 24 figg., 12

taw.

Volume XXV

I - CALEGARI G., 1989 - Le incisioni rupestri di Taouardei (Gao, Mali).

Problematica generale e repertorio iconografico, pp. 1-14, 9 figg., 24

taw.

li - PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia funzionale

e paleoecologia del rettile placodonte Psephoderma alpinum Meyer,

1858 .pp. 15-50, 20 figg., 9 taw.

Ili - CALDARA R., 1990 - Revisione Tassonomica delle specie paleartiche

del genere Tychius Germar (Coleoptera Curculionidae). pp. 51-218,

575 figg.

Volume XXVI

I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia, Placo-

dontia )./?/?. 1-21, 23 figg.

II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara preisto¬

rico: dati e interpretazioni./?/?. 25-556, 647 figg.

Ili - ANDRI E. e ROSSI F., 1993 - Genesi ed evoluzione di frangenti,

cinture, barriere ed atolli. Dalle stromatoliti alle comunità di scogliera

moderne, pp. 559-610, 49 figg., 1 tav.

Volume XXVII

I - PINNA G. and GH1SELIN M. edited by, 1996 - Biology as History. N.

1 . Systematic Biology as an Historical Science, pp. 1-133, 68 figs.

II - LEONARDI C. e SASSI D. a cura di, 1997 - Studi geobotanici ed en-

tomofaunistici nel Parco Regionale del Monte Barro, pp. 135-266, 122

figg., 23 tabb.

Volume XXVIII

I - BANFI E. & GALASSO G„ 1998 - La flora spontanea della città di

Milano alle soglie del terzo millennio e i suoi cambiamenti a partire dal

1700 .pp. 267-388, 71 figg., 30 tabb.

Volume XXIX

I - CALEGARI G., 1999 - L’arte rupestre dell’Eritrea. Repertorio ragionato

ed esegesi iconografica./?/?. 1-174, 268 figg.

Volume XXX

I - PEZZOTTA F. edited by, 2000 - Mineralogy and petrology of shallow

depth pegmatites. Paper from thè First International Workshop, pp. 1-

117, 30 figs., 19 tabs.

II - PARISI B., FRANCHINO A. & BERTI A. con la collaborazione di

POTENZA B. & RUBINI D., 2000 - La Società Italiana di Scienze

Naturali 1855 -2000. Percorsi storici./?/?. 1-163, 199 figg.

Ili - DE ANGELI A. & GARASSINO A., 2002 - Galatheid, chirostylid and

porcellanid decapods (Crustacea, Decapoda, Anomura) from thè Eoce¬

ne and Oligocene of Vicenza (N Italy )./?/?. 1-31, 21 figs., 9 pls.

Volume XXXI

I - NOSOTTI S. & RIEPPEL O., 2002 - The braincase of Placodus Agassiz,

1833 (Reptilia, Placodontia). pp. 1-18, 15 figs.

II - MARTORELLI G., 2002 - Monografia illustrata degli uccelli di rapina

in Italia. (1895). Riedizione a cura di Fausto Barbagli./?/?. [XX] 1-216,

[14] 46 figg., 4 tavv.

Ili - NOSOTTI S. & RIEPPEL O., 2003 - Eusanrosphargis dalsassoi n. gen.

n. sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano

(Lombardy, N Italy)./?/?. 1-33, 19 figs., 1 tab., 3 pls.

Volume XXXII

I - ALESSANDRELLO A., BRACCHI G. & RIOU B., 2004 - Polychaete,

sipunculan and enteropneust worms from thè Lower Callovian (Middle

Jurassic) of La Voulte-sur-Rhòne (Ardèche, France). pp. 1-16, 9 figs.,

1 pi.

II - RIEPPEL O. & HEAD J. J., 2004 - New specimens of thè fossil snake

genus Eupodophis Rage & Escuillié, from Cenomanian (Late Creta-

ceous) of Lebanon. pp. 1-26, 13 figs., 1 tab.

Ili - BRACCHI G. & ALESSANDRELLO A., 2005 - 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)./?/?. 1-48, 8 figs., 1 tab., 16 pls.

Volume XXXIII

I - BOESI A. & CARDI F. edited by, 2005 - Wildlife and plants in tradi-

tional and modem Tibet: conception, exploration and conservation. pp.

1-88, 30 figs., 9 tabs.

II - BANFI E., BRACCHI G„ GALASSO . & ROMANI E., 2005 - Agro-

stologia Piacentina, pp. 1-80, 7 figs., 1 tabs.

Ili - LIVI P. a cura di 2005 - I fondi speciali della Biblioteca del Museo

Civico di Storia Naturale di Milano. La raccolta di stampe antiche del

Centro Studi Archeologia Africana, pp. 1-250, 389 figs.

Volume XXXIV

I - GARASSINO A. & SCHWEIGERT G., 2006 - The Upper Jurassic

Solnhofen decapod crustacean fauna: review of thè types from old

descriptions. Pari I. Infraorders Astacidea, Thalassinidea and Palinura.

pp. 1-64, 12 figs., 20 pls.

I

I

Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali,

Museo Civico di Storia Naturale, Corso Venezia 55 -20121 Milano

Pubblicazione disponibile al cambio

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di Scienze Naturali

e del Museo Civico

Volume XXXIV - Fascicolo III di Storia Naturale di Milano

— - - — ■ —

A NEW SPECIES OF

PONTOSAURUS (SQUAMATA, PYTHONOMORPHA)

FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS

OF PYTHONOMORPHA

MILANO LUGLIO 2006

Elenco delle Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

Volume I

I - CORNALIA E., 1865 - Descrizione di una nuova specie dei genere

Felis: Fehs jacobita (Com.). 9 pp., 1 tav.

II - MAGNI-GRIFFI F., 1865 - Di una specie A'Hippolais nuova per l’Ita¬

lia. 6 pp., 1 tav.

Ili - GASTALDI B., 1865 - Sulla riescavazione dei bacini lacustri per opera

degli antichi ghiacciai. 30 pp., 2 figg-, 2 taw.

IV - SEGUENZA G., 1 865 - Paleontologia malacologica dei terreni terziarii

del distretto di Messina. 88 pp., 8 taw.

V - GIBELLI G., 1865 - Sugli organi riproduttori del genere Verrucaria,

16 pp., 1 tav.

VI - BEGGIATO F. S., 1865 - Antracoterio di Zovencedo e di Monteviale

nel Vicentino, 10 pp., 1 tav.

VII - COCCHI I., 1865 - Di alcuni resti umani e degli oggetti di umana

industria dei tempi preistorici raccolti in Toscana. 32 pp., 4 taw.

Vili - TARGIONI-TOZZETTI A., 1866 - Come sia fatto l’organo che fa

lume nella lucciola volante dell’Italia centrale (Luciola italica) e come

le fibre muscolari in questo ed altri Insetti ed Artropodi. 28 pp., 2 taw.

IX - MAGGI L., 1865 - Intorno al genere Aeolosoma. 18 pp., 2 taw.

X - CORNALLAE., 1865 - Sopra i caratteri microscopici offerti dalle Cantari¬

di e da altri Coleotteri facili a confondersi con esse. 40 pp., 4 taw.

Volume II

I - ISSEL A., 1 866 - 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 taw.

Ili - 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 taw.

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.

VII - COCCHI I., 1866 - L’uomo fossile nell’Italia centrale. 82 pp., 21 figg.,

4 taw.

Vili - GAROVAGLIO S., 1866 - Manzonia cantiana, novum Lichenum

Angiocarporum genus propositum atque descriptum. 8 pp., 1 tav.

IX - SEGUENZA G., 1867 - 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 taw.

Volume III

I - EMERY C., 1873 - Studii anatomici sulla Vipera Rechi. 16 pp., 1 tav.

II - GAROVAGLIO S., 1867 - Thelopsis, Beloraa, Weitenwebera et Limbo-

ria, quatuor Lichenum Angiocarporum genera recognita iconibusque

illustrata. 12 pp., 2 taw.

Ili - TARGIONI-TOZZETTI A., 1867 - Studii sulle Cocciniglie. 88 pp., 7 taw.

IV - CLAPARÈDE E. R. e PANCERI P., 1867 - Nota sopra un Alciopide

parassito della Cydippe densa Forsk. 8 pp. 1 taw.

V - GAROVAGLIO S., 1871 - De Pertusariis Europae mediae commenta-

tio. 40 pp., 4 taw.

Volume IV

I - D’ACHIARDI A., 1868 - Corallarj fossili del terreno nummulitico del-

l’ Alpi venete. Parte 11. 32 pp. 8 taw.

II - GAROVAGLIO S., 1868 - Octona Lichenum genera vel adhuc con¬

troversa, vel sedis prorsus incertae in systemate, novis descnptionibus

iconibusque 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 taw.

IV - (Non pubblicato).

V - MARINONI C., 1871 - Nuovi avanzi preistorici in Lombardia. 28 pp.,

3 figg., 2 taw.

NUOVA SERIE

Volume V

I - MARTORELLI G., 1895 - Monografia illustrata degli uccelli di rapina

in Italia. 216 pp., 46 figg., 4 taw.

Volume VI

I - DE ALESSANDRI G., 1897 - La pietra da cantoni di Rosignano e di

Vignale. Studi stratigrafici e paleontologici. 104 pp., 2 taw., 1 carta.

II - MARTORELLI G., 1898 - Le forme e le simmetrie delle macchie nel

piumaggio. Memoria ornitologica. 112 pp., 63 figg., 1 taw.

Ili - PAVESI R, 1901- L'abbate Spallanzani a Pavia. 68 pp., 14 figg., 1 tav.

Volume VII

1 - DE ALESSANDRI G., 1910 - Studi sui pesci triasici della Lombardia.

164 pp., 9 taw.

Volume Vili

1 - REPOSSI E., 1915 - La bassa Valle della Mera. Studi petrografici e

geologici. Parte I. pp. 1-46, 5 figg., 3 taw.

II - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi petrogra¬

fici e geologici. Parte II. pp. 47-186, 5 figg. 9 taw.

III - AIRAGIII 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 taw.

II - SERAG. L., 1920 - 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 prmcipali 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, pp. 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 taw.

III - SCORTECCI G., 1941- 1 recettori degli Agamidi. pp. 209-326, 80 figg.

Volume XI

I - GUIGLIA D., 1944 - Gli Sfecidi italiani del Museo di Milano (Hymen.).

pp. 1-44, 4 figg., 5 taw.

II-III - GIACOBINI 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 taw., 1 carta.

III - 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 taw., 1 carta.

II - PINNA G., 1963 - Ammoniti del Lias superiore (Toarciano) dell’Alpe

Turati (Erba, Como). Generi Mercaticeras, Pseudomercaticeras e Bro¬

di eia. pp. 65-98, 2 figg., 4 taw.

Ili - ZANZUCCHI G., 1963 - Le Ammoniti del Lias superiore (Toarciano)

di Entratico in Val Cavallina (Bergamasco orientale), pp. 99-146, 2

figg-- & tavv-

Volume XIV

I - VENZO S., 1965 - Rilevamento geologico dell’anfiteatro morenico

frontale 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.

Ili - DIENI I., MASSARI F. e MONTANARI L„ 1966 - Il Paleogene dei

dmtomi di Orosei (Sardegna), pp. 13-184, 5 figg., 8 taw.

Volume XV

I - C ARETTO 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 taw.

II - DIENI I. e MASSARI F., 1966 - Il Neogene e il Quaternario dei dintorni

di Orosei (Sardegna), pp. 89-142, 8 figg., 7 taw.

Ili - BARBIERI F„ IACCARINO S„ BARBIERI F. & PETRUCCI F., 1967

- Il Pliocene del Subappennino Piacentino-Parmense-Reggiano. pp.

143-188, 20 figg., 3 taw.

Volume XVI

I - C ARETTO P. G., 1967 - Studio morfologico con l’ausilio del metodo

statistico e nuova classificazione dei Gasteropodi pliocenici attribuibili

■d\Murex brandaris Linneo . pp. 1-60, 1 fig., 7 tabb., 10 taw.

II - SACCHI VIALLI G. e CANTALUPPI G., 1967-1 nuovi fossili di Goz¬

zano (Prealpi piemontesi)./?/?. 61-128, 30 figg., 8 taw.

Ili - PIGORINI B., 1967 - Aspetti sedimentologici del Mare Adriatico, pp.

129-200, 13 figg., 4 tabb. 7 taw.

Volume XVII

I - PINNA G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Alpe

Turati (Erba, Como). Famiglie Lytoceratidae, Nannolytoceratidae,

Hammatoceratidae (excl. Phymatoceratinae) Hildoceratidae (excl.

Hildoceratinae e Bouleiceratinae). pp. 1-70, 2 taw. n.t., 6 figg., 6 taw.

II - VENZO S. & PELOSIO G., 1968 - Nuova fauna a Ammonoidi del-

l’Anisico superiore di Lenna in Val Brembana (Bergamo), pp. 71-142,

5 figg., 11 taw.

Michael W. Caldwell

new species of Pontosaurus (Squamata, Pythonomorpha)

from thè Upper Cretaceous of Lebanon

and a phylogenetic analysis of Pythonomorpha

Volume XXXIV - Fascicolo III

Luglio 2006

Memorie della Società Italiana di Scienze Naturali

e del Museo Civico di Storia Naturale di Milano

INDEX

Introduction . Pag- ^

Pontosaurs, dolichosaurs and basai pythonomorphs:

an overview . Pag. 4

Systematic Palaeontology . Pag. 5

Osteological description . Pag.

State of preservation . Pag-

Skull . Pag-

Postcranial skeleton . Pag. 17

Axial skeleton . Pag. 1 7

Pectoral girdle and forelimb . Pag. 20

Pelvic girdle and rearlimb . Pag. 24

Squamation and tracheal rings . Pag. 28

Squamation . Pag- 28

Tracheal and ?bronchial? rings . Pag. 32

Phylogenetic analysis . Pag. 32

Methodology . Pag- 32

Results . Pag- 32

Character distributions . Pag. 34

Discussion . Pag. 35

Functional morphology . Pag. 35

Pontosaurus . Pag. 36

Acknowledgements . Pag. 37

References . Pag. 38

Appendix I . Pag. 40

Appendix II . Pag. 42

© 2006 Società Italiana di Scienze Naturali

Museo Civico di Storia Naturale

Corso Venezia, 55 - 20121 Milano

In copertina: Pontosaurus kornhuberi. Watercolour by 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 - Luglio 2006

ISSN 0376-2726

Michael W. Caldwell

A new species of Pontosaurus (Squamata, Pythonomorpha)

from thè Upper Cretaceous of Lebanon

and a phylogenetic analysis of Pythonomorpha

Abstract - A new fossil marine squamate from thè Upper Cretaceous (Cenomanian) of Lebanon is described and its phylogenetic relationships

analyzed. The taxon represents a new species of Pontosaurus , previously known from only thè type species and only one specimen, designated

Pontosaurus lesinensis (Kornhuber, 1873). The new taxon is very well preserved including thè squamation and a complete tail numbering 163 caudal

vertebrae. New and important features of thè braincase of pontosaurs are highlighted: 1) thè supraoccipital position and its overlap of thè parietal; 2)

thè alar process of prootics elongate contact with parietal; 3) thè reduction of thè paroccipital process. A phylogenetic analysis of pythonomorphs, uti-

lizing these new characters in a data matrix of 77 characters and 10 terminal taxa, finds nine shortest trees with a tree length of 1 19 steps (C.I. = 0.773;

HI = 0.226). The nine shortest trees vary only in thè sistergroup relationships of Aphanizocnemus libanensis within non-mosasauroid pythonomorphs,

and thè ingroup relationships of ophidian pythonomorphs. For example, cladogram one presents thè following sistergroup structure: ((Aigialosauridae,

Mosasauridae) (Dolichosauridae ( Aphanizocnemus libanensis (( Pontosaurus lesinensis, Pontosaurus kornhuberi) ( Adriosaurus suessi (Pachyophis

woodwardi (Pachyrhachis problematicus, Serpentes))))))). In thè context of thè hypothesized phylogeny presented in this study, thè functional mor-

phology of pontosaurs is discusseci, highlighting tail and limb function in relationship to locomotory capability.

Key words - Squamates, pythonomorphs, marine lizards, Cretaceous, Lebanon, phylogeny.

Riassunto - Una nuova specie di Pontosaurus (Squamata, Pythonomorpha) dal Cretacico superiore del Libano e una analisi filogenetica dei

Pythonomorpha.

Viene descritto un nuovo squamato fossile marino e ne vengono analizzate le relazioni filogenetiche. Il taxon rappresenta una nuova specie di

Pontosaurus, genere di cui era precedentemente nota solamente la specie tipo, con un singolo esemplare, denominata Pontosaurus lesinensis

(Kornhuber, 1873). Il nuovo esemplare qui descritto è molto ben conservato: la coda di 163 vertebre è completa e sono presenti resti delle squame.

Vengono messe in luce nuove e importanti caratteristiche della scatola cranica: 1) la posizione del sopraoccipitale e la sua sovrapposizione al parieta¬

le; 2) l’esteso contatto del processo alare dei prootici con il parietale; 3) la riduzione del processo paraoccipitale. Una analisi filogenetica dei pitono-

morfi che utilizza questi nuovi caratteri in una matrice di 77 caratteri e 10 taxa terminali produce nove alberi con una lunghezza di 1 19 steps (CI =

0.773; HI = 0.226). Questi nove alberi variano unicamente nelle relazioni filogenetiche di Aphanizocnemus libanensis entro i pitonomorfi non-mosa-

sauroidi, e nelle relazioni filogenetiche interne dei pitonomorfi ofidi. Ad esempio il primo cladogramma presenta la seguente articolazione dei succes¬

sivi sistergroups: ((Aigialosauridae, Mosasauridae) (Dolichosauridae ( Aphanizocnemus libanensis ((Pontosaurus lesinensis, Pontosaurus kornhuberi )

(Adriosaurus suessi (Pachyophis woodwardi (Pachyrhachis problematicus, Serpentes)))))))). La morfologia funzionale dei pontosauri viene discussa

nel contesto della filogenesi ipotizzata, evidenziando Timportante funzione locomotoria della coda e degli arti.

Parole chiave - Squamati, pitonomorfi, rettili marini, Cretacico, Libano, filogenesi.

INTRODUCTION

Over thè course of their known history, encompassing

more than 130 million years, squamate reptiles evolved a

wide and surprising array of anatomies as thè various

clades radiated and adapted into both terrestrial and

aquatic environments. For example, limb reduction to

complete limblessness, usually associated with extreme

increases in body length, is observed in almost all lineag-

es of living squamates (Greer, 1991; Caldwell, 2003) and

certainly characterizes thè nearly 3,000 species of snakes,

which are arboreal, terrestrial, fossorial, cryptic, and

aquatic. Modern squamates also show an amazing range

in size from thè incredibly large komodo dragons to thè

smallest known living amniotes Sphaerodactylus ariasae,

a Caribbean gecko, which is only 16 millimeters long

(Hedges & Thomas, 2001).

The list of modern squamate adaptations and anatom-

ical innovations allowing them to colonize varied envi¬

ronments is extensive with but two exceptions - thè air

and thè sea. Among more than 7,000 species of living

squamates, none have evolved powered flight though

some do glide, and only 59 species have evolved and

adapted to life in aquatic marine environments: one

species of iguana, 50 species of true sea snakes, 5 species

of sea kraits, and 3 species of file snakes.

Interestingly, while these living squamates show

remarkable adaptations to life in marine environments, thè

true acme of squamate evolution in marine environments

is not found among modern species, but rather among fos¬

sil squamates. It is thè extinct pythonomorphs, i.e., thè

dolichosaurs, pontosaurs, aigialosaurs, mosasaurs, and

4

MICHAEL W. CALDWELL

early snakes, which truly became “aquatic” in every sense

nearly 100 million years ago. This aspect of squamate evo-

lution and evolutionary history is very poorly detailed in

thè literature purporting to evaluate aspects of squamate

evolution. For example, thè recent and excellent treatise on

lizard diversity and evolution by Pianka & Vitt (2003: 260)

only uses thè word “mosasaur” once in more than three

hundred pages, and never even mentions thè existence of

aigialosaurs, dolichosaurs, pontosaurs, adriosaurs or coni-

asaurs. As a result, there is also no discussion of thè extent

of pythonomorph evolution and adaptation to marine envi-

ronments. This knowledge vacuum can only be rectified

by thè presentation of synthetic works, fusing phylogeny,

function, and anatomy, that detail these missing pieces of

squamate evolutionary history.

This study presents a description and phylogenetic

analysis of a new species of ancient marine squamate,

assigned to thè genus Pontosaurus, that was found in 95

million year old rocks (Upper Cretaceous) in Lebanon.

This new specimen is remarkably complete, including thè

preservation of soft tissues, and thus permits a unusual

degree of “biological” interpretation (PI. 1). The genus

Pontosaurus was previously based on only one specimen,

GBA 1873/4/2 (GBA, Austrian Geological Survey, Wien,

Austria), thè type species, Pontosaurus lesinensis

(Kornhuber, 1873). Pontosaurus lesinensis was found in

rocks outcropping on thè island of Hvar, Croatia (Pierce

& Caldwell, 2004) that are considered to be chrono-

equivalents of those in Lebanon that produced thè new

species described here.

The new specimen was collected from a quarry in thè

Valley of Al Gabour near Al Nammoura, 10 km southeast

of Hadjula, Lebanon (Dal Sasso & Pinna, 1997; Dal

Sasso & Renesto, 1999). Previous to this work. Dal Sasso

& Pinna (1997) had described thè first known lizard from

thè marine sediments of Lebanon, Aphanizocnemus liba-

nensis, which is considered to be Middle Cenomanian in

age and was collected at a third quarry near Hakel,

Lebanon. The specimen described here is therefore thè

second known lizard from thè platy limestones of

Lebanon and is a very different animai compared to A.

libanensis.

Recent work by Caldwell & Dal Sasso (2004)

described thè remarkably well-preserved squamation of

thè new lizard, a feature not usually preserved in verte¬

brate fossils and certainly not for squamates. In this study

thè squamation will be reviewed as part of thè overall

description of thè specimen, but thè focus will be on thè

osteology. Characters obtained from this study will be

added to a modified version of thè character matrix of

Pierce & Caldwell (2004) for thè purpose of constructing

a phylogenetic hypothesis of all pythonomorphs.

PONTOSAURS, DOLICHOSAURS AND BASAL PYTHONOMORPHS: AN OVERVIEW

In 1873, Kornhuber described a new genus and species

of Cretaceous lizard from two separate specimens pre¬

served on platy limestone slabs that had come from quar-

ries outcropping on thè island of Hvar, Croatia, known in

Italian as Isola di Lesina. Kornhuber’s piate “A” preserved

thè posterior portion of thè skeleton of a small reptile; this

piate made its way to Vienna, Austria, by 1869.

Kornhuber’s second specimen, piate “B”, arrived in Vienna

in 1870 and preserved thè anterior portion, including thè

head, neck and forelimbs, of a second small reptile.

Kornhuber’s comparison of thè two specimens led him to

conclude they were thè same thing, if not thè same individ¬

uai, and he therefore assigned them to thè same species,

Hydrosaurus lesinensis ; thè genus Hydrosaurus (Wagler,

1830) is now recognized as thè junior synonym of thè

extant monitor lizard genus, Varanus (Merrem, 1 820).

In 1892, Gorjanovic-Kramberger renamed Hydrosaurus

lesinensis as Pontosaurus lesinensis (Kramberger, 1892)

based on his assessment that it was not similar enough to

thè extant Varanus ( =Hydrosaurus ) to warrant assignment

to that genus. In thè act of recognizing yet another fossil

Adriatic lizard, including Aigialosaurus dalmatici is

(Kramberger, 1892), Adriosaurus suessi (Seeley, 1881),

and Acteosaurus tomassinii (Meyer, 1860), Kramberger

created thè Family Aigialosauridae to include all thè fossil

forms.

Nopcsa (1903) reevaluated thè systematic position of P

lesinensis in a study that included all of Kramberger’s ( 1 892)

taxa, as well as Dolichosaurus longicollis (Owen, 1850),

Opetiosaurus bucchichi (Kornhuber, 1901), Carsosaurus

marchesetti (Kornhuber, 1893), and Mesoleptos zendrinii

(Comalia, 1851). Nopcsa (1903) identified common features

shared by Pontosaurus , Acteosaurus, Adriosaurus and

Dolichosaurus, and assigned them all to thè family

Dolichosauridae (Gervais, 1852), while Aigialosaurus ,

Opetiosaurus, Carsosaurus, and Mesoleptos were assigned

to thè family Aigialosauridae.

Since Nopcsa (1903), except for brief mentions in com¬

parative studies (Nopcsa, 1908, 1923; Calligaris, 1987/1988;

Rieppel, 1988; Dal Sasso & Pinna, 1997; Dal Sasso &

Renesto, 1999), Pontosaurus had never been reexamined in

any detail until thè study of Pierce & Caldwell (2004). The

latter study attempted to assess Nopcsa’s (1903) assignment

of Pontosaurus to thè Dolichosauridae based primarily on

recent analyses of existing specimens of Adriosaurus,

Acteosaurus, Dolichosaurus and Coniasaurus (Caldwell,

1999a, 1999b; Caldwell, 2000; Caldwell & Cooper, 1999;

Lee & Caldwell, 2000; Caldwell & Lee, 2004).

Owen (1850) described two monotypic genera of

marine lizards, Dolichosaurus longicollis and Coniasaurus

crassidens, based on a number of specimens collected

from thè Lower Chalk (Cenomanian; Upper Cretaceous) of

southeast England. Dolichosaurus was diagnosed by a

uniquely high number of cervical and dorsal vertebrae

while unique tooth characters diagnosed Coniasaurus',

Owen did not identify any characters linking these two

taxa. Nopcsa (1908) restudied Owen’s specimens and

added several new specimens to thè list of known coni-

A NEW SPECIES OF PONTOSAURUS (SQUAMATA, PYTHONOMORPHA) FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS OF PYTHONOMORPHA

5

asaurs and dolichosaurs. Unfortunately for Coniasaurus,

Owen (1850) had misidentified thè tooth-bearing element

of thè type specimen (a maxilla) as a dentary leaving

Nopcsa (1908) to build on this error by first, not finding

thè true type, and second, by identifying a pair of well-pre-

served dentaries as Owen’s type specimen (see Caldwell &

Cooper [1999] for thè venue of thè holotype and a

redescription of thè type material). Caldwell’s (1999a)

description of Coniasaurus gracilodens highlighted sever-

al notable differences with C. crassidens such as tooth

shape, maxillary tooth number, and robustness of thè max¬

illa (thè maxilla of C. crassidens is much more elongate

than that of C. gracilodens ). Interestingly, all species

assigned to Coniasaurus are known only from disarticulat-

ed skulls and isolated vertebral elements. In contrast, thè

type and referred specimens of Dolichosaurus longicollis

are articulated postcranial skeletons with only one very

fragmentary skull and no preserved teeth (Caldwell, 2000).

Without consideration of pontosaurs, adriosaurs, or

acteosaurs, coniasaurs and dolichosaurs have been thought

to be nested within basai mosasauroids (mosasaurs and

aigialosaurs) since Nopcsa, (1908), or recently, since

Caldwell’s (1999b) hypothesis that they are thè sistergroup

to mosasauroids. The difficulty in resolving thè

Coniasaurus - Dolichosaurus problem is that both taxa

are represented by non-comparable data: Dolichosaurus

is known from complete postcrania with little or no skull

material, while both species of Coniasaurus are known

from skulls and only a small number of disarticulated

vertebrae.

Nopcsa’s (1903) suggestion that pontosaurs, adriosaurs,

SYSTEMATIC

Reptilia Linnaeus, 1758

Squamata Oppel, 1811

Pythonomorpha Cope, 1 869

Genus Pontosaurus Kramberger, 1 892

Type Species: Hydrosaurus lesinensis

Komhuber, 1873

Pontosaurus lesinensis (Kornhuber, 1873)

Holotypus: GBA 1873/4/2 (GBA, Austrian Geological

Survey, Wien, Austria): articulated cranial and postcranial

remains of one individuai; postcranium includes 12 artic¬

ulated cervical vertebrae, 28 dorsal vertebrae, shoulder

girdles, forelimbs, fragments of femur, and pelvic girdle.

Locus typicum: Hvar Island (= Isola di Lesina),

43°10’N, 16°30’E, Croatia; upper Cenomanian-upper

Turonian (Upper Cretaceous).

Generic Diagnosis: Long, slender lizard possessing

unique supraoccipital-parietal articulation, with supraoc-

cipital resting on top of and forming w-shaped suture

with parietal; elongation of all postdentary bones; 10-12

cervical vertebrae; hypapophyses with large, unfused

peduncles on C2-C10; 26-28 dorsal vertebrae; fused

scapulocoracoid with primary coracoid emargination.

acteosaurs, coniasaurs, and dolichosaurs might all be

assigned to thè Dolichosauridae, in association with recent

studies proposing a sistergroup relationship between all

these taxa and thè mosasauroids (Caldwell, 1999b; Lee &

Caldwell, 2000), highlight thè phylogenetic importance of

these animals in understanding thè evolution of Mesozoic

marine squamates. Recent work by Caldwell & Lee (1997),

Lee & Caldwell (1998, 2000), and Caldwell (1999b), all of

which examined thè larger problem of squamate phylogeny,

found Serpentes to be nested within a squamate clade

including dolichosaurids and mosasauroids, a group

referred to as thè Pythonomorpha (see Cope’s [1869] orig¬

inai diagnosis of this group as contrasted with Lee &

Caldwell’s [2000] slightly modified definition).

To date, thè one hundred and fifty-five years of investi-

gation on Cenomanian marine squamates has achieved thè

consensus view that thè fossil forms (i.e., pontosaurs,

adriosaurs, acteosaurs, coniasaurs, and dolichosaurs) are

closely related to each other; there is also a reasonable

degree of consensus that these squamates share a common

ancestor with mosasaurids and aigialosaurids. However, as

to thè phylogenetic position of this larger group within

squamates, and of snakes with any one group of squamates,

there is no common agreement. For thè purposes of this

study, I will consider pontosaurs, adriosaurs, acteosaurs,

coniasaurs, and dolichosaurs as informai generic groupings

assigned to thè Pythonomorpha; thè Dolichosauridae, thè

conventional taxon for all these taxa, will be treated as

including only Dolichosaurus and Coniasaurus.

Pontosaurus kornhuberi sp. nov.

(Figs. 1,2, 3A-B, 4-18)

Synonomy: cf. Pontosaurus sp. Caldwell & Dal Sasso,

2004.

Derivatio nominis: named in honour of Prof. A.

Kornhuber in recognition of his contributions to thè study

of pythonomorph squamates.

Holotypus: MSNM V3662: complete, articulated

skeleton of one individuai (1,037 mm in length), includ¬

ing trachea, bronchi and squamation, preserved on platy

limestone slab, housed in thè Museo di Storia Naturale di

Milano.

Diagnosis: Maxilla with large medially inflected pre-

maxillary process; jugal with distinct posteroventral

tuberosity; parietal with broad fiat, medially expanded

dorsal table extending onto parietal rami; frontal margin

emarginated with no distinct nasal processes; 10-11 cer¬

vical vertebrae; 26 dorsal vertebrae; 2 pygal vertebrae;

163 caudal vertebrae; non-imbricated scales on cheek,

imbricated scales on remainder of body.

Locus typicum: Quarry in thè Valley of Al Gabour

near Al Nammoura, 10 km southeast of Hadjula, Lebanon

(Dal Sasso & Pinna, 1997; Dal Sasso & Renesto, 1999).

6

MICHAEL W. CALDWELL

Fig. 1 - Holotype, Pontosaurus kornhuberi , MSNM V3662. A) Interpretative drawing; B) photo. Scale bar = 10 cm.

A NEW SPECIES OF PONTOSAURUS (SQUAMATA, PYTHONOMORPHA) FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS OF PYTHONOMORPHA

7

OSTEOLOGICAL DESCRIPTION

State of preservation

MSNM V3662 possesses a long (46 mm long), thin

skull (18.6 mm wide) with large orbits (Figs. 1,2,3 A-B)

and an elongated postorbitai region. The skull is flattened

and preserved in dorsal view, but was clearly compressed

from thè left as thè left mandible is exposed in lateral view

while thè right has rotated medially underneath thè derma-

tocranium; only thè mediai aspect of thè right mandible is

visible to thè right of thè parietal. Likewise, thè left maxil-

la has shifted into lateral view, and is broken away; thè right

maxilla is preserved in dorsal view, but is crushed down-

wards so that thè ascending process is compressed onto thè

maxillary shelf. The neck bears at least 10 and probably 1 1

cervical vertebrae with ribs, and a trachea extending to thè

third dorsal rib. The limbs are well developed, and thè forc¬

are both well preserved and in articulation. There are 26

pachyostotic trunk vertebrae and ribs; pachyostosis is not

evident in thè sacrai or caudal skeleton. The tail is excep-

tionally long (680 mm) and possesses 163 caudal vertebrae;

thè number of caudals is reduced by two from thè 1 65 as

reported by Caldwell & Dal Sasso (2004) due to thè recog-

nition in this study of two probable pygal vertebrae. The

haemal spines are almost twice as long as thè neural spines

and articulate with distinct haemapophyses; transverse

processes disappear at thè 8^ caudal vertebra. Detailed

preservation in this fine-grained limestone has preserved

thè squamation both as molds, casts, and in many places,

thè originai material included tonai variation.

Skull

Fig. 2 - Pontosaurus kornhuberi, MSNM V3662, detail of thè skull.

limb (65 mm long) is reduced in size relative to thè rear-

limb (98 mm long); thè right forelimb is not well exposed

but thè left is well preserved and articulated. The rearlimbs

The skull is represented by thè premaxilla, right max¬

illa and partial left maxilla, fragments of thè septomaxil-

lae, vomers, and possibly nasals, partial right and left pre-

frontals, complete right and left jugals, thè left postor¬

bitai, postfrontal, and squamosal, and fragments of thè

right-side series of thè same, a well preserved frontal, and

parietal, thè supraoccipital, a well -preserved left exoccip-

ital-opisthotic and fragments of thè right, probable left

and right prootics, a left quadrate with fragments of thè

right, a well preserved left dentary with teeth, a well pre¬

served left surangular with a poorly exposed right suran-

gular, left and right articulars (Figs. 2, and 3 A-B), and

perhaps thè quadrate ramus of thè right pterygoid.

Premaxilla

The premaxilla is a single bone and is well preserved

though broken into small fragments along thè length of

thè premaxillary bar (Fig. 4 A-B). The anterior tooth-

bearing portion of thè premaxilla (Fig. 4 C-D) is a small

median element that flares right and left to support two

teeth on each side of thè sagittal piane (Figs. 2, 3, and 4).

At its anterior tip, it is broad and fiat with a small space

equivalent to thè width of one tooth base separating thè

right and left pairs of teeth; thè anterior extent of thè bone

is posteriorly deepened just enough to support thè alveoli

for thè four teeth and immediately becomes constricted

posterior to this point in order to form a long and thin nar-

ial bar similar to that of Pontosaurus lesinensis (Fig. 3 C-

D) (Pierce & Caldwell, 2004).

The premaxilla and its internarial bar is much better

preserved in MSNM V3662 than that of thè type of

Pontosaurus lesinensis (Fig. 3 A-D) and shows an impor¬

tai feature shared in common with Coniasaurus gracilo-

dens Caldwell, 1999. On thè right side of thè premaxilla

in MSNM V3662 there is a small, laterally positioned

fossa or small platform that lies below thè level of thè

internarial bar and immediately posterior to thè tooth

hearing portion of thè bone (Fig. 4 C-D). Immediately lat¬

eral, and stili in proximate articulation is a matching,

8

MICHAEL W. CALDWELL

Fig. 3 - Skulls of Pontosaurus kornhuberi MSNM V3662 (A-B) and Pontosaurus lesinensis GBA 1873/4/2 (C-D). A) Dorsal surface of skull; B) line draw-

ing ot sanie; C) dorsal surtace of skull; D) line drawing of same. Abbreviations: a, atlas intercentrum; an, atlas neural arch; ax, axis; co, coronoid; d, den-

tary; ec, ectopterygoid; epi, epipterygoid; f, frontal; j, jugal; mx, maxilla; na, nasal; p, parietal; p-r, parietal ramus; pf, postfrontal; pmx, premaxilla; pof,

postorbito frontal; po, postorbitai; popr, paroccipital process of thè exoccipital-opisthotic; prf, prefrontal; prò, prootic; pt, pterygoid; q, quadrate; rap, retroar-

ticular process; smx-vo-p, septomaxilla and vomeropalatine; soc, supraoccipital; sq, squamosal; st, supratemporal; sa, surangular.

A NEW SPECIES OF PONTOSAURUS (SQUAMATA, PYTHONOMORPHA) FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS OF PYTHONOMORPHA

9

Fig. 4 - Snout of Pontosaurus kornhuberi, MSNM V3662. A) OverView of palatal elements viewed through external narial opening; B) line drawing

with palatal and snout elements illustrated in color (premaxilla = red; septomaxilla and vomeropalatine = blue; prefrontal = yellow; possible nasal frag-

ment = grey); C) photo detail of premaxilla showing right and lefit maxillae, premaxillary bar, and well preserved premaxillary process of right max-

illa; D) line drawing of same with premaxilla highlighted in red. Abbreviations: see Fig. 3.

10

MICHAEL W. CALDWELL

anteromedially directed hook-shaped process extending

from thè maxillary shelf of thè right maxilla. The sidewall

of thè right maxilla is compressed on top of thè maxillary

shelf but thè separation of thè transverse shelf and verti-

cal sidewall are stili easily detected (Fig. 4 C-D).

The morphology and position of this anatomical fea-

ture is identical to that of Coniasaurus gracilodens

(Caldwell, 1999a, figs. 6 B, and 7 B). As was suggested

by Caldwell (1999a), this process, likely a synapomorphy

of C. gracilodens and P. kornhuberi, suggests that thè

very small premaxilla of thè latter and by extension thè

probably equal sized premaxilla of thè former, would

have been mobile around these paired processes. The

morphology of this anatomical feature hints at an unsus-

pected degree of premaxillary mobility in these small

pythonomorph reptiles. The maxilla was not firmly

sutured to thè premaxilla, thè premaxilla was long, and

there is no evidence of a firm suture with thè frontal at thè

posterior tip of thè internarmi bar. Similar snout kinesis is

observed in modern snakes with thè notable difference

being thè significantly shorter narial bar (Frazetta, 1970).

The internarial bar of Pontosaurus kornhuberi appears

to bear a tiny notch at thè posteriormost tip (Fig. 4 A-B);

presumably, this notch contacted some portion of thè

internarial process of thè frontal. It is not clear how firm¬

ly sutured this contact was, but it appears to have been

rather loose. In contrast to P. kornhuberi , thè internarial

bar of P lesinensis is sandwiched between a pair of nasal

bones situated along its lateral borders that appear to

restrict thè premaxillary contribution to thè external nari¬

al opening and seem to prevent its contact with thè frontal

bone, a scenario consistent with all squamates except

mosasauroids (Fig. 3 C-D; Pierce & Caldwell, 2004). If

nasals were present in P kornhuberi, they are not well

represented in MSNM V3662 by anything more than one

possible sliver of bone lying in thè left narial opening

(Fig. 4 A-B). It is also possible that this fragment is a por¬

tion of thè palatine and not thè nasal. Further support for

thè absence of nasal bones is thè different frontal margin

morphology of P kornhuberi as compared to P. lesinensis

(thè former is an effaced “w”-shape while thè latter is a

“w-w”-shape hearing two small nasal processes parasagit-

tal to thè internarial process [Fig. 3 C-D]).

Maxilla

The left maxilla is very poorly preserved and will not be

described here. The right is much better preserved (Figs. 2,

3 A-B, 4) even though it is only visible in dorsal view; an

accurate tooth count is therefore problematic. As described

previously, thè anterior tip of thè maxilla is rounded and

rises gently into a shallow but vertically oriented lateral wall

that frames thè margin of thè narial opening (Fig. 4 A-B).

Mediai to thè sidewall of thè maxilla and near thè anterior

tip of thè bone, is a well-developed premaxillary process

that extends medially and anteriorly towards a fossa or

recess on thè premaxilla; thè maxillary shelf, beginning

immediately posterior to thè premaxillary process, bears a

scalloped margin lateral to its contact with thè vomers. In

dorsal view thè anterior two-thirds of thè maxilla shows an

identical morphology to thè maxilla of Coniasaurus gracilo¬

dens (Caldwell, 1999a). The difference between thè maxil-

lae of P. kornhuberi and C. gracilodens is thè height and

extent of thè ascending process of thè maxilla in thè latter

taxon. Pontosaurus kornhuberi possesses a low ascending

process, similar to P. lesinensis (Fig. 3 A-D) and a fairly long

anteriorly directed prefrontal that blocks any articulation

between thè frontal and maxilla. In P. lesinensis thè ascend¬

ing process is a long, low sloping feature of thè maxilla that

rises to form only about one half of thè lateral margin of thè

narial opening (Fig. 3 C-D); a similar condition also charac-

terizes P. kornhuberi except that thè maxillary contribution

to thè narial opening is less, comprising about two fifths of

thè margin (Figs. 3 A-B, 4 A-B).

The posterior process of thè maxilla extends below thè

orbit only far enough to contact thè anterior tip of thè

jugal. However, this distance appears to be equal to about

one half of thè orbit diameter; a similar condition is

observed in all mosasauroid pythonomorphs (Russell,

1967). Due to preservation and orientation of thè element,

it is not possible to determine how far posteriorly, and

perhaps how far beneath thè orbit, thè dentition extended.

In Pontosaurus lesinensis (Pierce & Caldwell, 2004),

Coniasaurus gracilodens (Caldwell, 1999a) and C. cras-

sidens (Caldwell & Cooper, 1999), thè dentition termi-

nates at thè anterior margin of thè orbit, as it does in other

mosasauroids; it seems likely that thè same condition

exists in P. kornhuberi.

Although a maxillary tooth count is not possible,

some aspects of thè maxillary dentition are preserved on

thè left side of thè skull and are worth describing. Most of

thè anterior teeth have been crushed and broken but some

are well enough preserved so that their generai outline

can be observed: sharp slightly recurved piercing teeth set

within a mass of ossified tissue as in other mosasauroids

(Russell, 1967; Carroll & deBraga, 1992; deBraga &

Carroll, 1993; Caldwell et al., 2002); fluting similar to

that observed in Pontosaurus lesinensis is not visible.

Septomaxilla

The septomaxillae are likely preserved in Pontosaurus

kornhuberi, but it is not possible to differentiate thè mor¬

phology of either thè left or right element due to dorsal-

ventral compaction of thè bone on top of vomeropalatines

(Fig. 4 A-B). Comparing thè morphology of thè anterior

portion of thè compressed septomaxilla/vomeropalatine

mass to thè probable septomaxilla of Coniasaurus

gracilodens (Caldwell, 1999a, fig. 9), where thè latter is

shaped like an obtuse triangle with a tali, median crest

paralleling thè internarial bar indicates that thè septomax¬

illa of Pontosaurus kornhuberi was similarly shaped and

anteriorly located. However, it also must be remembered

that this is not a certain identification but rather a discus-

sion of thè generai similarities between these two taxa; by

comparison, there are no preserved remnants of thè sep¬

tomaxillae in Pontosaurus lesinensis (Fig. 3 C-D).

Vomeropalatines

As with thè septomaxilla, it is not possible, due to

A NEW SPECIES OF PONTOSAURUS (SQUAMATA, PYTHONOMORPHA) FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS OF PYTHONOMORPHA

11

preservational factors, to determine whether or not thè

vomers and palatines were separate bones. As it has been

shown that thè two elements are fused in mosasaurs

(Russell, 1967) and Coniasaurus gracilodens (Caldwell,

1999a, figs. 10, 11), these two elements are discussed

here as though they were a single fused structure (thè ele¬

ments are not distinguishable in Pontosaurus lesinensis

[see Pierce & Caldwell, 2004]).

The vomeropalatines of MSNM V3662 are elongate,

strap-like elements that parallel thè sagittal piane but do

not appear to meet at thè midiine (Fig. 4 A-B). The poste-

rior extent of thè bone appears to be beneath thè frontal

(thè palatine ramus), but there is some uncertainty as

exposed and this statement must be made only tentatively.

Anteriorly, at a point equivalent to thè anterior termination

of thè prefrontal, thè vomeropalatine extends a lateral

process to contact thè maxilla (maxillary process of thè

palatine); at this point thè vomeropalatine continues ante¬

riorly following thè margin of thè maxillary shelf to thè

mediai divergence of that shelf where it forms thè premax-

illary process (Fig. 4 A-B). The tips of thè vomeropalatine

appear to diverge away from thè midi ine as they converge

towards thè premaxillary process. Details such as thè posi-

tion of Jacobson’s Organ, or thè aperture of thè mediai

palatine sinus, are not visible as preserved.

Prefrontal

The prefrontal is an anteriorly elongate element with

thè anterior process extending at least three-fifths of thè

length of thè narial opening along thè mediai surface of

thè side wall of thè maxilla. Posterodorsally, thè pre¬

frontal contacts thè frontal along about one quarter of thè

length of thè frontal and then descends ventrally to con¬

tact thè anterior tip of thè jugal and thè posterior process

of thè maxilla. Overall, thè exposed surface of thè right

prefrontal, as preserved, resembles thè sail of a tali sail-

boat (Figs. 2, 3 A-B, 4 A-B). The articulation with thè

maxilla is an important and characteristic feature of pon-

tosaurs and is shared in common by both species: thè

suture is long and low and slopes gently towards thè nar¬

ial opening. The result is that there is no ascending

process of thè maxilla, creating a similar profile to that

observed in many extant snakes; this feature contrasts

sharply with that described by Caldwell (1999a) for

Coniasaurus gracilodens where thè maxilla has a well-

developed ascending process. The difference between

Pontosaurus lesinensis and P. kornhuberi is that in thè

former thè maxilla and prefrontal comprise equal por-

tions of thè margin of thè narial opening, whereas, as

noted for thè latter, thè prefrontal is three fifths of thè

margin of thè opening.

Nasals

As noted previously, it is not clear from thè specimen

(Fig. 4 A-B) as to thè presence or absence of thè nasal bones.

This is in direct contrast to thè relatively well-preserved

nasals of Pontosaurus lesinensis , where thè nasals are thin,

fiat and paired, (Pierce & Caldwell, 2004) (Fig. 3 C-D).

Lacrimai

It is not possible to identify thè presence or absence of

a lacrimai in Pontosaurus kornhuberi as thè area on either

side of thè skull, ventral to thè prefrontals, is heavily

crushed and deformed (Figs. 2, 3 A-B, 4 A-B). As was

noted by Pierce & Caldwell (2004), Kornhuber (1873)

had reported a trace of thè right lacrimai posterodorsal to

thè maxilla and anterior to thè jugal, but such an element

was not visible to thè former authors. The massive size of

thè prefrontal would suggest, if analogous to snakes, that

thè lacrimai is absent in both species of Pontosaurus as it

is in snakes, Pachyrhachis (Lee & Caldwell, 1998) and

many other squamate groups (Estes et al ., 1988).

Jugal

Both thè right and left jugal are well preserved in

Pontosaurus kornhuberi (Figs. 2, 3 A-B, 5 A-C). The ante¬

rior edge of thè jugal arch extends forward to meet thè pre¬

frontal anteriorly and thè maxilla ventrally and may well

have extended anteriorly past thè orbit as in other

mosasauroids (deBraga & Carroll, 1993); this observation

is contra thè condition reported in P lesinensis by Pierce &

Caldwell (2004) except that thè anteriormost tip of thè jugal

is not visible and so may well have overlapped thè maxilla,

thus extending anterior to thè orbit.

A second difference is also noted between thè two

species: in P. kornhuberi thè two rami of thè jugal form a

distinct angle at thè posteroventral corner of thè orbit pro-

ducing a well-developed posteroventral tuberosity. In

contrast, in P. lesinensis thè jugal curves gradually in an

anterior direction to contact thè lateral process of thè pos-

torbital. In all other respects, thè jugal of Pontosaurus

lesinensis is of roughly similar size and shape to that of P

kornhuberi (Fig. 3 A-D).

Frontal

The frontal is a long, fiat element that comprises

approximately one-third of thè total length of thè skull

(Figs. 2, 3 A-B, 5 A-B). As is common to pontosaurs and

coniasaurs thè outline in dorsal view resembles an hour-

glass that is broadly expanded at both its anterior and pos¬

terior ends and is constricted above thè orbits (Caldwell,

1999a; Pierce & Caldwell, 2004); this configuration is

very similar to that of modera macroteiids (Estes et al.,

1988). In Pontosaurus kornhuberi, thè anterior expansion

of thè frontal is only marginally more narrow than thè

posterior expansion at thè frontal-parietal suture; this

contrasts with thè condition in P lesinensis where thè

anterior expansion is noticeably more narrow than thè

expansion at thè frontal-parietal suture, and with

Coniasaurus gracilodens (Caldwell, 1999a, figs. 4, 5)

where thè posterior expansion is twice thè width of thè

anterior expansion.

In Pontosaurus kornhuberi, thè anterior border of thè

frontal has only three processes (Figs. 2, 3 A-B, 4 A-B): one

large sagittal process and two equal-sized large lateral

processes. In contrast, thè frontal of P lesinensis has five

processes: one large sagittal process, two large lateral

processes, and two small parasagittal processes (Fig. 3 C-

12

MICHAEL W. CALDWELL

D). In both species, thè external edge of thè lateral process-

es articulates with thè prefrontals, while thè internai rim

forms thè posterior margin of thè external narial opening.

Pierce & Caldwell (2004) noted that thè nasals of P.

lesinensis were clasped by thè unpaired sagittal, and paired

parasagittal, processes. The absence of thè paired parasagit-

tal processes in P. kornhuberi may well relate to thè absence

of identifiable nasal bones. The sagittal process of

Pontosaurus kornhuberi is smooth as it is in P. lesinensis ;

in contrast, thè sagittal process of Coniasaurus gracilodens

bears a low but clearly defined sagittal crest that extends

posteriorly about one quarter of thè length of thè frontal.

At its midpoint, thè constricted frontal forms thè dor-

sal margin of thè orbit. The prefrontal and postor-

bitofrontal do not meet above thè orbits as they do in

higher mosasauroids (Russell, 1967), but are widely sep-

arated over thè orbit as they are in Pontosaurus lesinensis

(Fig. 3 C-D) and Coniasaurus gracilodens (Caldwell,

1999a, figs. 4, 5).

The posterior border of thè frontal describes an obtusely

angled w-shape along its margin with thè parietal; this mar¬

gin represents a gap of considerable size and is not interpret-

ed as an artifact of preservation, but rather is very well pre-

served. It is possible that this morphology represents thè

adult conditimi and that there was a high degree of mesoki-

nesis in pontosaurs. It is also possible that ffontal-parietal

ossification was delayed in this animai, either that it was a

younger individuai or that in fact dermatocranial ossifica¬

tion was delayed in Pontosaur kornhuberi. In comparison,

thè frontal-parietal margin in Pontosaurus lesinensis is rela-

tively simple and straight with thè two bones fitting togeth-

er very tightly (Fig. 3 C-D).

Parietal

The parietal is a single element composing approxi-

mately one-third of thè total skull length (Figs. 2, 3 A-B,

5 C, 6 A-D). In generai terms, thè suture with thè frontal

is long and describes an obtuse “w”-shaped outline with

large parietal lappets lying below thè frontal. Along thè

lateral edge of thè frontal-parietal suture, thè anterolater-

al projection of thè parietal is broadly overlapped by thè

postorbitofrontal. Laterally and ventrally, thè parietal is

broadly expanded to form a long and deep decensus pari-

etalis (Figs. 3 A-B, 5 C, 6 A) which rises slightly at its

midpoint at thè suturai contact with an element identified

here as thè prootic. The dorsal surface of thè parietal is

not developed into a long thin crest as in P lesinensis

(Fig. 3 C-D), but rather is broad, fiat table-like surface.

The parietal foramen is not as large as that of P. lesinen¬

sis; it is also an equivalent distance posterior to thè pari-

etal-frontal suture in both species.

The prootic margin of thè parietal also appears to con¬

tact thè margin of thè supraoccipital; thè latter margin is

broad and wide in its contact with thè parietal and is in thè

sanie piane; thè supraoccipital does not He beneath thè pari¬

etal as in lizards but is in thè sanie piane and articulates

along its anterior margin as in snakes. There is also an inter-

digitating and overlapping suturai contact between thè

supraoccipital and parietal where thè supraoccipital over-

lies thè parietal on thè midiine in large triangular shaped

depression (Fig. 6 A-D). This same condition is observed in

Pontosaurus lesinensis (Fig. 3 C-D). The differences

between P. kornhuberi and P. lesinensis are preservational:

in thè former, thè overlapping portion of thè supraoccipital

is broken away leaving a fractured surface on thè supraoc¬

cipital, whereas in P. lesinensis thè small sagittal-crest like

structure of thè supraoccipital is stili in place.

The parietal ramus for thè suspensorium is well devel¬

oped though only well preserved on thè left side of thè

skull, where it contacts a small supratemporal element

and thè small, underlying paroccipital process of thè

exoccipital-opisthotic. The parietal ramus of Pontosaurus

kornhuberi is broken away and displaced vertically from

its position at thè posterior edge of thè decensus parietal-

is. In its proper articulation, thè parietal ramus would

have been oriented obliquely and would have formed a

continuous surface with thè decensus parietalis as in P.

lesinensis (Fig. 3 C-D).

Postorbitofrontal

The postfrontal and postorbitai of Pontosaurus kornhu¬

beri appear to have fused to form a single, multi-rami ele¬

ment, thè postorbitofrontal; thè element is very well pre¬

served on thè left side of thè skull, but very poorly pre¬

served on thè right. All descriptions given here are of thè

left side of thè skull. Close inspection indicates thè pres-

ence of an unidentified element immediately anterior to thè

left postorbitofrontal (Figs. 3 A-B, 5 A-C) that is considered

here to be thè left postfrontal. Squamates show a perplex-

ing and problematic degree of plasticity in thè presence,

absence, or fusion, of thè elements comprising thè anterior

portions of thè supratemporal arcade. For example, some

groups possess only a postorbitai (snakes), while others

possess both postfrontals and postorbitals (e.g.,

Pontosaurus lesinensis [Fig. 3 C-D]), while others, such as

species assigned to thè varanid genus, Varanus, and P.

kornhuberi , possess a single, apparently fused, postor¬

bitofrontal element. Some possess supernumerary ele¬

ments, such as pythons, which have added a supraorbital

bone. Assuming that thè anterior element is not a true post¬

frontal, thè remaining description will focus on thè mor¬

phology of thè left postorbitofrontal.

The left postorbitofrontal is a quadri-radiate element

with four distinct rami contacting four very different

components of thè dermatocranium. The anterior or

frontal ramus is a moderately long and thin spline of bone

that begins at roughly thè frontal-parietal suture and

clasps thè lateral margin of thè frontal to a position rough¬

ly near thè beginning of thè supraorbital constriction

(Figs. 2, 3 A-B, 5 A-C). The mediai or parietal ramus of

thè postorbitofrontal is very short and medially directed.

It clasps thè margin of thè parietal table just posterior to

thè frontal-parietal suture; there is a short section of thè

postorbitofrontal, between thè frontal and parietal rami,

that firmly abuts thè lateral wing of thè parietal immedi¬

ately posterior to thè frontal-parietal suture. The lateral or

jugal ramus of thè postorbitofrontal projects laterally only

a short distance; this ramus is thickened posteriorly with

A NEW SPECIES OF PONTOSAURUS (SQUAMATA. PYTHONOMORPHA) FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS OF PYTHONOMORPHA

13

Fig. 5 - Mid-dorsal surface of thè dermatocranium of Pontosaurus kornhuberi, MSNM V3662. A) Photo detail of lcft postorbital-frontal-parietal; B)

line drawing of same; C) oblique view of left side of skull showing configuration of decensus parietalis and contact with Ieft prootic. Abbreviations:

see Fig. 3.

14

MICHAEL W. CALDWELL

I ig. 6 - Posterodorsal surface of dermatocranium of Pontosaurus kornhuberi, MSNM V3662. A) OverView of parietal; B) detail of supraoccipital and

overlapping, interdigitating suturai contact onto parietal; C) detail of parietal ramus, squamosal, exoccipital, supraoccipital; D) line drawing of ‘C\ E)

left quadrate. F) line drawing of same. Abbreviations: see Fig. 3.

A NEW SPECIES OF PONTOSAURUS (SQUAMATA, PYTHONOMORPHA) FROM THE UPPER CRETACEOUS OF LEBANON

AND A PHYLOGENETIC ANALYSIS OF PYTHONOMORPHA

15

! its anteriormost edge beginning parallel to thè piane of

thè frontal-parietal suture. It is not clear if some portion

of thè jugal ramus was directed ventrally; however, it is

possible that thè ascending portion of thè jugal contacted

a small facet on thè ventral surface of thè jugal ramus

similar to some mosasaurs (Russell, 1967) and many

squamates (Estes et al., 1988). The posterior or squamos-

al ramus of thè postorbitofrontal is elongate and thin

(Figs. 5 A-C, 6 A-C) extending posteriorly to a point

equal to thè contact of thè parietal and supraoccipital. It

overlaps thè squamosal laterally and dorsally in an

interfingering suture. The distance between thè postor¬

bitofrontal bar and thè parietal crest is rather far even

though thè supratemporal fenestra appears to be relative-

ly small anteriorly (Figs. 5 A-C, 6 A-C).

Postfrontal?

It is not clear if thè postfrontal is present as a separate

element in Pontosaurus kornhuberi even though it is

clearly separate in P. lesinensis (Fig. 3 C-D). The postor¬

bitofrontal, described previously, shows thè typical squa¬

mate morphology of a fused postorbitai ramus and body

(posterior ramus contacting thè squamosal and framing

thè supratemporal fenestra, main body contacting thè

jugal) and postfrontal ramus (anterior process clasping

thè frontal-parietal suture and extending anterolaterally

along thè frontal). However, there is an extra element in P

kornhuberi that lies anterior and lateral to thè anterior

ramus of postorbitofrontal thè identity of which is uncer-

tain (Figs. 2, 3 A-B, 5 A-C). The element is indicated with

a “?” on Fig. 3 B. It is possible that it is a fragment of thè

pterygoid, or perhaps is a displaced palpebrai bone, but as

most of thè skull is very well articulated, thè latter possi-

bility is considered remote. The element bears a striking

resemblance to, and is in a similar position to, thè post¬

frontal of P. lesinensis which is a large boomerang-shaped

element (Fig. 3 C-D) that clasps both thè frontal and pari¬

etal along its mediai border and attaches to, but does not

fuse with, thè postorbitai along its posterolateral surface.

The fusion of thè postfrontal and postorbitai in form-

ing thè postorbitofrontal is not a well-understood process,

nor is it common among squamates. It is also clear ffom

a survey of modern squamates that there is great deal of

plasticity in thè formation and fusion of thè dermatocra-

nial elements of thè supratemporal and postorbitai

arcades (see Estes et al., 1988). Non-scolecophidian

snakes are considered to possess only a postorbitai with

thè exception of Dinilysia patagonica which also possess-

es a discrete postfrontal (Caldwell & Albino, 2002);

amphisbaenians and dibamids possess neither element;

Varanus possesses a single postorbitofrontal element

while Lanthanotus and Heloderma have only a post¬

frontal with thè postorbitai having been lost; anguids typ-

ically have both a postfrontal and postorbitai, as do most

other scincomorphs; gekkotans are treated as possessing

a postorbitofrontal even though thè posterior ramus is

missing and iguanians show both a single element and

paired element condition.

Squamosal

The squamosal is a thin band of bone that forms thè pos¬

terolateral border of thè supratemporal fenestra (Figs. 3 A-B,

5 C, 6 C-D). At thè midpoint of thè supratemporal fenestra,

thè squamosal articulates with thè ramus of thè postor¬

bitofrontal; it underlies that element in a long, clasping artic-

ulation (Fig. 6 C-D). At its posterior tip, thè squamosal con-

tacts thè quadrate laterally and thè supratemporal medially.

The element is slightly bent laterally at its posterior contact

with thè supratemporal and quadrate. A similar morphology

is observed in Pontosaurus lesinensis (Fig. 3 C-D).

Supratemporal

The supratemporal is a small wedge of bone that as

preserved, appears to articulate with thè squamosal,

quadrate and parietal ramus. Presumably it also contacted

thè paroccipital process of exoccipital-opisthotic; howev¬

er, this is not clear as preserved and it is important to note

that thè paroccipital process appears to be quite short.

Supraoccipital

The supraoccipital of Pontosaurus kornhuberi (Figs.