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MEMORIE
della Società Italiana
di Scienze Naturali
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Volume XXXV - Fascicolo I di Storia Naturale di Milano
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U1®SITY
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
CATALOG AND BIBLIOGRAPHY
OF THE FOSSIL
STOMATOPODA AND
DECAPODA
FROM ITALY
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MILANO NOVEMBRE 2006
Elenco delle Memorie della Società Italiana di Scienze Naturali
e del Museo Civico di Storia Naturale di Milano
Volume 1
I - CORNALI A E., 1865 - Descrizione di una nuova specie dei genere
Felis: Felis jacobita (Corn.). 9 pp., 1 tav.
II - MAGNI-GRIFFI F., 1865 - Di una specie d 'Hippolais nuova per l’Ita¬
lia. 6 pp., 1 tav.
Ili - GASTALDI B., 1865 - Sulla riescavazione dei bacini lacustri per opera
degli antichi ghiacciai. 30 pp., 2 figg.. 2 tavv.
IV - SEGUENZA G„ 1865 - Paleontologia malacologica dei terreni terziarii
del distretto di Messina. 88 pp., 8 tavv.
V - GIBELLI G„ 1865 - Sugli organi riproduttori del genere Vermcaria,
16 pp., 1 tav.
VI - BEGGIATO F. S., 1865 - Antracoterio di Zovencedo e di Monteviale
nel Vicentino, 10 pp., 1 tav.
VII - COCCHI I., 1865 - Di alcuni resti umani e degli oggetti di umana
industria dei tempi preistorici raccolti in Toscana. 32 pp., 4 tavv.
Vili - TARGIONI-TOZZETTI A., 1866 - Come sia fatto l’organo che fa
lume nella lucciola volante dell’Italia centrale (Luciola italica) e come
le fibre muscolari in questo ed altri Insetti ed Artropodi. 28 pp., 2 tavv.
IX - MAGGI L., 1865 - Intorno al genere Aeolosoma. 18 pp., 2 taw.
X - CORNALI A 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., 1866 - Quelques considérations sur l'origine des bassins
lacustres, àpropos des sondages du Lac de Come. 12 pp., 8 tavv.
Ili - MOLON F., 1867 - Sulla flora terziaria delle Prealpi venete. 140 pp.
IV - D’ACHIARDI A., 1866 - Corallarj fossili del terreno nummulitico
delle Alpi venete. 54 pp., 5 tavv.
V - COCCHI I., 1866 - Sulla geologia dell’alta Valle di Magra. 18 pp., 1 tav.
VI - SEGUENZA G., 1866 - Sulle importanti relazioni paleontologiche di
talune rocce cretacee della Calabria con alcuni terreni di Sicilia e del¬
l’Africa settentrionale. 18 pp., 1 tav.
VII - COCCHI I., 1 866 - L’uomo fossile nell’Italia centrale. 82 pp., 21 figg.,
4 taw.
Vili - G ARO VAGLIO S., 1866 - Manzonia cantiana, novum Lichemim
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 tavv.
Volume III
I - EMERY C., 1873 - Studii anatomici sulla Vipera Redii. 16 pp., 1 tav.
II - GAROVAGLIO S., 1867 - Thelopsis, Belonia, Weiterrwebera 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 tavv.
IV - CLAPARÈDE E. R. e PANCERI P„ 1867 - Nota sopra un Alciopide
parassito della Cydippe densa Forsk. 8 pp. 1 tavv.
V - GAROVAGLIO S., 1871 - De Pertusariis 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 vel adhuc con¬
troversa, vel sedis prorsus incerlae in systemate, novis descriptionibus
iconibusque accuratissimis illustrata. 18 pp., 2 tavv.
Ili - MARINONI C., 1868 - Le abitazioni lacustri e gli avanzi di umana
industria in Lombardia. 66 pp., 5 figg., 7 tavv.
IV - (Non pubblicato).
V - MARINONI C., 1871 - Nuovi avanzi preistorici in Lombardia. 28 pp.,
3 figg., 2 tavv.
NUOVA SERIE
Volume V
1 - MARTORELLI G., 1895 - Monografia illustrata degli uccelli di rapina
in Italia. 216 pp., 46 figg., 4 tavv.
Volume VI
I - DE ALESSANDRI G., 1897 - La pietra da cantoni di Rosignano e di
Vignale. Studi stratigrafici e paleontologici. 104 pp., 2 tavv, 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 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.
1 64 pp., 9 taw.
Volume Vili
1 - REPOSSI E., 1915 - La bassa Valle della Mera. Studi petrografia e
geologici. Parte I .pp. 1-46, 5 figg., 3 tavv.
II - REPOSSI E., 1916 ( 1917) - La bassa Valle della Mera. Studi petrogra¬
fia 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, pp.
1-164, 7 figg. 2 taw.
II - SERA G. L., 1920 - Sui rapporti della conformazione della base del cra¬
nio colle fonrie 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, 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.
Ili - SCORTECCI G., 1941- 1 recettori degli Agamidi. pp. 209-326, 80 figg.
Volume XI
1 - GUIGLIA D.. 1944 - Gli Sfecidi italiani del Museo di Milano (Hymen.).
pp. 1-44, 4 figg., 5 tavv.
II-I1I - 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), pp. 1-70, 4 figg. 6 taw.
I - VENZO S., 1957 - Rilevamento geologico dell’anfiteatro morenico del
Garda. Parte I: Tratto occidentale Gardone-Desenzano. pp. 71-140, 14
figg., 6 tavv, 1 carta.
Ili - VIALLI V., 1959 - Ammoniti sinemuriane del Monte Albenza (Berga¬
mo), pp. 141-188, 2 figg., 5 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 tavv, 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 tavv.
III - ZANZUCCHI G., 1963 - Le Ammoniti del Lias superiore (Toarciano)
di Entratico in Val Cavallina (Bergamasco orientale), pp. 99-146, 2
figg., 8 taw.
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 Dactvlioceratidae. pp. 83-136, 4 taw.
Ili - DIENI 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., 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- Di ENI I. e MASSARI F., 1966 - Il Neogene e il Quaternario dei dintorni
di Orosei (Sardegna), pp. 89-142, 8 figg., 7 taw.
III - BARBIERI F„ IACCAR1NO S„ BARBIERI F. & PETRUCCI F„ 1967
- 11 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 tavv
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, Nannolyloceratidae,
Hammatoceratidae (excl. Phvmatoceratinae) Hildoceratidae (excl.
flildoceratinae e Bouleiceratinae). pp. 1-70, 2 tavv. n.t., 6 figg.. 6 tavv
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.
Antonio De Angeli & Alessandro Garassino
Associazione Amici del Museo Zannato di Montecchio Maggiore
Museo Civico di Storia Naturale di Milano
Catalog and bibliography of thè fossil
Stomatopoda and Decapoda from Italy
Volume XXXV - Fascicolo I
Novembre 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: from left to right, Titanocarcinus aculeatus Busulini, Tessier & Visentin, 1 984 (Reconstruction A. De Angeli),
Rosenfeldia triasica Garassino, Teruzzi & Dalla Vecchia, 1996 (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 - Novembre 2006
ISSN 0376-2726
Antonio De Angeli & Alessandro Garassino
Catalog and bibliography of thè fossil
Stomatopoda and Decapoda from Italy
INTRODUCTION
The rich Italian decapod faunas, reported from thè
Triassic to Pleistocene, have been thè subject of many
studies by Italian and foreign researchers for over two
centuries. Important studies were carried out by famous
palaeontologists, such as Ristori, Crema, A. Sismonda,
E. Sismonda, Bittner, A. Milne Edwards, Reuss, and
Lovisato, who pioneered thè systematic study of Italian
decapod crustaceans between thè middle of thè nineteenth
century and thè beginning of thè last century. Unfortu-
nately, not many studies were carried out in thè first half
of thè twentieth century following this initial effort. In thè
last thirty years, thè discovery of rich decapod faunas in
many outcrops reported in Piemonte, Lombardia, Veneto,
Friuli-Venezia Giulia, and Campania has given a new
impulse to thè carcinological studies. The numerous
systematic papers, published in these last years by Itali-
an researchers and collaborators of thè Museo Civico
di Storia Naturale di Milano and of thè Museo Civico
“G. Zannato” di Montecchio Maggiore (Vicenza), have
resulted in an increase in thè knowledge of Mesozoic and
Cenozoic decapod crustaceans of Italy by thè descriptions
of new genera and species of stomatopods, anomurans,
macrurans and brachyurans.
The main purposes of this catalog are essentially three:
1 - To locate thè Italian Museum or University in
which thè nineteenth century collections are housed in
order to give an updated check list of thè holotypes.
2 - To give an updated check list of thè carcinological
studies under way today in Italy.
3 - To provide a valid work instrument for thè rease-
archers to supplement “Treatise on Invertebrate Paleon-
tology”, published by Glaessner (1969).
Finally, we note that thè research of thè originai speci-
mens from Sardegna, studied by Meneghini, Lòrenthey,
Lovisato, Comaschi Caria, and Marras & Ventura; from
Puglia, studied by Varola, Bonfiglio & Donadeo, and
Ristori; and from Piemonte, studied by Allasinaz was
very difficult because of thè catalogue numbers and, in
some cases, thè entire collections are missing. Moreover,
we note that many specimens housed in Museo dei Fos¬
sili di Bolca (Verona) and Museo Civico “G. Zannato”
di Montecchio Maggiore (Vicenza) are reported with thè
acronym I.G. (Inventario Generale dello Stato).
The systematic arrangement used in this catalog
follows thè classifìcation proposed by Martin & Davis
(2001).
ABBREVIATIONS FOR INSTITUTIONS HOLDING SPECIMENS
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
SYSTEMATIC CATALOG
Order Stomatopoda Latreille, 1817
Superfamily Gonodactyloidea Giesbrecht, 1910
Family Pseudosquillidae Manning, 1977
Pseudosquilla Dana, 1852
Type species: Squilla ciliata Fabricius, 1787.
Stratigraphic range: Eocene - Recent.
Pseudosquilla berica De Angeli & Messina, 1996
1996 - Pseudosquilla berica De Angeli & Messina; p. 6, Text-fig. 2,
PI. 1 (fig. 1 a-c)
2000 - Pseudosquilla berica De Angeli & Messina in Hof; p. 35
2001 - Pseudosquilla berica De Angeli & Messina in De Angeli &
Beschin; p. 40, Text-fig. 35
2004 - Pseudosquilla berica De Angeli & Messina in Schram &
Muller; p. 76
2006 - Pseudosquilla berica De Angeli & Messina in De Angeli &
Rossi; p. 86, PI. 2 (fig. 5)
Holotype: MCZ 1547 (I.G. 284625).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli & Messina (1996) reported one
specimen (MCZ 1547) from Perarolo (Vicenza).
Superfamily Lysiosquilloidea Giesbrecht, 1910
Family Lysiosquillidae Giesbrecht, 1910
Lysiosquilla Dana, 1852
Type species: Squilla scabricauda Lamarck, 1818.
Stratigraphic range: Cretaceous - Recent.
Lysiosquilla antiqua (Munster, 1842)
1842 - Squilla antiqua Munster; p. 76, PI. 9 (fig. 11)
1915 - Squilla antiqua Munster in Fabiani; p. 284
1975 Lysiosquilla antiqua (Munster) in Secretan; p. 371, Text-figs.
23-32, Pls. 26-33
1975 - Lysiosquilla antiqua var. minor Secretan; p. 378, Text-fig. 33,
Pls. 34-37, nov syn.
1996 - Lysiosquilla antiqua (Munster) in De Angeli & Messina; p. 9
2004 - Lysiosquilla antiqua (Munster) in Schram & Muller; p. 90
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Munster ( 1 842) reported one specimen from
Monte Bolca (Verona) (repository and catalogue number
unknown); Secretan (1975) reported 35 specimens from
thè sanie quarry (MSNV x, 03, 50, 50bis, 51, 5 Ibis, 52,
53, 54, 56, 58, 57, from 59 to 68, from 70 to 84, 86, 88, 89,
ZI, Z2, B7, B8, B55, B55bis, FI, F3, F4, F6, Cr4, Cr5;
MGPD 10923, 12546, 12549).
Note: Schram & Muller (2004) considered L. antiqua
var. minor Secretan, 1975, as synonym with L. antiqua
(Munster, 1842).
Lysiosquilla messinai De Angeli, 1997
1997 - Lysiosquilla messinae De Angeli; p. 24, Tex-fig. 1
2001 - Lysiosquilla messinae De Angeli in De Angeli & Beschin; p. 40
2004 - Lysiosquilla messinae De Angeli in Schram & Mailer; p. 91
2006 - Lysiosquilla messinai De Angeli in De Angeli & Rossi; p. 86,
PI- 2 (fig. 2)
Holotype: MCZ 1546 (I.G. 284624).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli (1997) reported one specimen
(MCZ 1546) from Perarolo (Vicenza).
Superfamily Squilloidea Latreille, 1802
Family Squillidae Latreille, 1802
Squilla Fabricius, 1787
Type species: Cancer mantis Linnaeus, 1758.
Stratigraphic range: (?Cretaceous) Eocene
Recent.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
5
Squilla miocenica Lovisato, 1 894
1894 - Squilla miocenica Lovisato; p. 207, Text-fìgs. 1-3
1949 - Squilla miocenica Lovisato in Via Boada; p. 2, Text-fig. 1
1956 - Squilla miocenica Lovisato in Comaschi Caria; p. 288
1996 - Squilla miocenica Lovisato in De Angeli & Messina; p. 9
1998 - Squilla miocenica Lovisato in Hof; p. 1570
2004 - Squilla miocenica Lovisato in Schram & Mùller; p. 199
Holotype: unknown.
Stratigraphic range: middle Miocene (Badenian).
Occurrence: Sardegna.
Material: Lovisato (1894) reported three specimens
from Cagliari (repository and catalogne number unknown).
Note: this species is also reported from thè upper
Miocene (Tortonian) of Montjuich (Spain).
Indeterminates
Genere e specie indeterminati
2005 - Genere e specie indeterminati in Beschin, De Angeli, Checchi &
! Zarantonello; p. 26, Text-fig. 18, PI. 4 (fig. 7)
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported one specimen
(MCZ 1554) from Grola di Comedo Vicentino (Vicenza).
Order Decapoda Latreille, 1 802
Suborder Dendrobranchiata Bate, 1888
Superfamily Penaeoidea Rafìnesque-Schmaltz, 1815
t Family Aegeridae Burkenroad, 1963
Aeger Mtinster, 1839
Type species: Macrourites tipularius Schlotheim, 1822.
Stratigraphic range: Upper Triassic (Camian) -
Upper Cretaceous (Cenomanian).
Aeger elongatus Garassino & Temzzi, 1990
1990 - Aeger elongatus Garassino & Teruzzi; p. 118, Text-fig. 24
Holotype: MSNM Ì8772.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino & Temzzi (1990) reported two
specimens (MSNM Ì8771, Ì8772) from Osteno (Como).
Aeger foersteri Garassino & Temzzi, 1990
1990 -Aeger foersteri Garassino & Teruzzi; p. 106, Text-figs. 1, 10-12
Holotype: MSNM Ì8749.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino & Temzzi (1990) reported seven
specimens (MSNM from Ì8742 to Ì8746, Ì8748, Ì8749)
from Osteno (Como).
Aeger macropus Garassino & Temzzi, 1990
1990 - Aeger macropus Garassino & Teruzzi; p. 117, Text-fig. 23
Holotype: MSNM Ì8770.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino & Temzzi (1990) reported one
specimen (MSNM Ì8770) from Osteno (Como).
Aeger muensteri Garassino & Temzzi, 1990
1990 - Aeger muensteri Garassino & Teruzzi; p. 109, Text-figs 2-3,
13-14
Holotype: MSNM Ì8752.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino & Temzzi (1990) reported three
specimens (MSNM Ì8750, Ì8751, Ì8752) from Osteno
(Como).
Aeger robustus Garassino & Temzzi, 1 990
1990 - Aeger robustus Garassino & Teruzzi; p. 112, Text-figs. 4,
15-16, 18-21
Holotype: MSNM Ì8759.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino & Temzzi (1990) reported ten
specimens (MSNM from Ì8753 to Ì8762) from Osteno
(Como).
Aeger rostrospinatus Garassino & Temzzi, 1990
1990 - Aeger rostrospinatus Garassino & Teruzzi; p. 1 14, Text-figs.
5-8, 22
Holotype: MSNM Ì8768.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino & Temzzi (1990) reported six spec¬
imens (MSNM from Ì8763 to Ì8769) from Osteno (Como).
Aeger sp.
1974 -Aeger sp. in Pinna; p. 21, PI. 1 (fig. 4)
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Pinna (1974) reported one specimen
(MSNB 3160) from Cene (Bergamo).
Family Penaeidae Rafìnesque-Schmaltz, 1815
Antrimpos Munster, 1839
Type species: Antrimpos speciosus Munster, 1839.
Stratigraphic range: Upper Triassic (Camian)
Upper Jurassic (Portlandian).
Antrimpos noricus Pinna, 1974
1974 - Antrimpos noricus Pinna; p. 14, Text-figs. 4-8, Pls. 2-11
1976 - Antrimpos noricus Pinna in Pinna; p. 34, PI. 1 (figs. 1-2), PI. 3
(hg- 2)
r
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
6
1990 - Antrimpos sp. in Dalla Vecchia; p. 137, Text-fig. 37
1991 - Antrimpos sp. in Dalla Vecchia; p. 26
1996 - Antrimpos noricus Pinna in Garassino, Teruzzi & Dalla Vec¬
chia; p. 30
Holotype: MSNB 3380.
Stratigraphic range; Upper Triassic (Norian).
Occurrence: Lombardia, Friuli Venezia-Giulia.
Material: Pinna (1974) reported 149 specimens (thè
best specimens are: MSNB 3110, 3111, 3122, 3130, 3133,
3 1 34, 3 1 36, 3 1 37, 3380; MSNM Ì4480, Ì4485) from Cene
(Bergamo); Pinna (1976) reported 34 specimens (thè best
specimens are: MSNBS 2771, 2773, 2780, 2932, 2978)
from Rest (Brescia); Garassino et al. (1996) reported 11
specimens (MFSN 1421, 1536 b-c, 1655, 16096, 16101,
16109, 16142 a-b, 16166, 1722 a-b, 1731, 13720) from
Rio Seazza (Udine).
Antrimpos sp.
2001 - Antrimpos sp. in Garassino & Teruzzi; p. 189, Text-fig. 1
Stratigraphic range: Lower Jurassic (Toarcian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (2001) reported one
specimen (MSNM il 0852) from Sogno (Bergamo).
Dusa Miinster, 1839
Type species: Dusa monocera Miinster, 1839.
Stratigraphic range: Upper Triassic (Norian) - Upper
Jurassic (Portlandian).
Dusa longipes (Pinna, 1974)
1974 - Palaeodusa longipes Pinna; p. 21, Text-figs. 9-10, PI. 1 (fig. 5),
PI. 12 (fig. 1), PI. 13 (figs. 1-3)
1976 - Palaeodusa longipes Pinna in Pinna; p. 36, Text-fig. 1, PI. 2
(figs. 1-2)
1990 - Dusa denticulata Miinster in Dalla Vecchia; p. 122, Text-
fig. 21, Pls. 31-35
1991 - Dusa denticulata Miinster in Dalla Vecchia; p. 22, Text-
figs. 15-17
1996 - Dusa longipes (Pinna) in Garassino, Teruzzi & Dalla Vecchia;
p. 24, Text-figs. 2-5, 11-15
Holotype: MSNB 3422.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia, Friuli-Venezia Giulia,
Lazio.
Material: Pinna (1974) reported 43 specimens from
Cene (Bergamo), (MSNB, catalogue number unknown);
Pinna (1976) reported three specimens (MSNBS 2930,
2937, 3012) from Rest (Brescia); Garassino et al. (1996)
reported 58 specimens (MFSN 992, 1416, 1417, 1418,
1420, 1427, 1431, 1433, 1434, 1439, 1442, 1446, 1449,
1469, 1474, 1475, 1491, 1493, 1494, 1509, 1554, 1588,
1590, 1592, 1604, 1680, 1688, 1697, 1698, 1703, 1715,
1730, 1893, 1895, 1951, 6441, 13717, 13722, 13725,
13804, 13808, 13809, 15381, 16094, 16097, 16098,
16099, 16100, 16103, 16105, 16106, 16107, 16167,
16168, 16169, 16385, 18455) from Rio Seazza, Rio
Forchiar, Forni di Sopra, Rio di Donna, and Rio Secco
(Udine).
Note: Dalla Vecchia (1991) reported some specimens
(thè best specimens are: MFSN 1607, 6441) ascribing
them to Dusa denticulata Miinster, 1839. Probably these
specimens could be ascribed to Dusa longipes.
Dusa cfr. D. longipes (Pinna, 1974)
1993 - Palaeodusa cfr. P. longipes Pinna in Garassino & Teruzzi;
p. 1 1, Text-fig. 16
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported two spec¬
imens (MSNB 7737, 8314) from Ponte Giurino (Bergamo).
Dusa sp.
1993 - Dusa sp. in Dalla Vecchia; p. 62, Text-figs. 3-4
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lazio.
Material: Dalla Vecchia (1993) reported 11 specimens
(MFSN from 15100 to 15111) from Filettino (Lazio).
Longichela Garassino & Teruzzi, 1993
Type species: Longichela orobica Garassino & Te¬
ruzzi, 1993.
Stratigraphic range: Upper Triassic (Norian).
Longichela orobica Garassino & Teruzzi, 1993
1993 - Longichela orobica Garassino & Teruzzi; p. 5, Text-figs. 3-8,
PI. 1 (figs. 1-2)
Holotype: MSNM Ì10738.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported 154
specimens (thè best specimens are: MSNM il 0739,
il 0744, i 1 075 1 , i 1 0752; MSNB 7725, 7748, 7751, 7760)
from Ponte Giurino (Bergamo).
Longichela cfr. L. orobica Garassino & Teruzzi, 1993
1993 - Longichela cfr. L. orobica in Garassino & Teruzzi; p. 8, Text-
figs. 9-11, PI. 1 (fig. 3)
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported two
specimens (MSNB 8198, 8199-8200) from Ponte Giurino
(Bergamo).
Micropenaeus Bravi & Garassino, 1998
Type species: Micropenaeus tenuirostris Bravi &
Garassino, 1998.
Stratigraphic range: Lower Cretaceous (Albian).
Micropenaeus tenuirostris Bravi & Garassino, 1 998
1998a - Micropenaeus tenuirostris Bravi & Garassino; p. 152, Text-
fig. 23
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
7
Holotype: M 21833.
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998a) reported
one specimen (M 21833) from Pietraroia (Beneven¬
to).
Penaeus Fabricius, 1798
Type species: by subsequent designation of Latreille,
1810, Penaeus monodon Fabricius, 1798.
Stratigraphic range: Lower Cretaceous (Hauterivian-
Barremian) - Recent.
Penaeus bolcensis Secretan, 1975
1975 - Penaeus bolcensis Secretan; p. 327, Text-figs. 4-6,
Pls. 2-4
1996 -Penaeus bolcensis Secretan in Garassino & Teruzzi; p. 4
1999 - Penaeus bolcensis Secretan in Beschin & Garassino; p. 194
Holotype: MSNV 100.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported one specimen
(MSNV 100) from Monte Bolca (Verona).
Penaeus cornappensis Garassino, 1998
1998 - Penaeus cornappensis Garassino; p. 60, Text-fig. 1
Holotype: MFSN 21536.
Stratigraphic range: Lower Cretaceous (Barremian -
Aptian).
Occurrence: Friuli-Venezia Giulia.
Material: Garassino (1998) reported four specimens
(MFSN 21535, 21536, 21542, 21543) from Torrente Cor-
nappo (Udine).
Penaeus obtusus Secretan, 1975
1975 - Penaeus obtusus Secretan; p. 330, Text-figs. 7-9, Pls. 7-10
1996 - Penaeus obtusus Secretan in Garassino & Teruzzi; p. 4
1999 - Penaeus obtusus Secretan in Beschin & Garassino; p. 194
Holotype: MSNV 106.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported one specimen
(MSNV 106) from Monte Bolca (Verona).
Penaeus sorbinii Beschin & Garassino, 1999
1999 - Penaeus sorbinii Beschin & Garassino; p. 201, Text-
figs. 8-10
2001 - Penaeus sorbinii Beschin & Garassino in De Angeli & Beschin;
p. 10
Holotype: MSNM Ì24505.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin & Garassino (1999) reported three
specimens (MSNM i 1 2633, Ì24505; MCZ 1583) from
Salcedo (Vicenza).
Penaeus vanzii Beschin & Garassino, 1 999
1999 - Penaeus vanzii Beschin & Garassino; p. 197, Text-figs. 4-7
2001 - Penaeus vanzii Beschin & Garassino in De Angeli & Beschin;
p. 10, Text-fig. 4
Holotype: MSNM Ì24501.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin & Garassino (1999) reported 26
specimens (MSNM Ì8408, i9 1 64, i9 1 65-Ì9 1 66, i9 1 67-
i9 1 69, from Ì9173 to Ì9176, Ì12634, Ì13593, from Ì24501
to Ì24504; MCZ from 1580 to 1582 and from 1604 to
1612) from Valle del Ponte (Vicenza).
Penaeus vernassensis Garassino & Teruzzi, 1995
1995 - Penaeus vernassensis Garassino & Teruzzi; p. 80, Text-fig. 2
Holotype: MFSN 17052.
Stratigraphic range: Lower Cretaceous (Hauterivian -
Barremian).
Occurrence: Friuli-Venezia Giulia.
Material: Garassino & Teruzzi (1995) reported three
specimens (MFSN 4308 a-b, 4310, 17052) from Vemasso
(Udine).
Pseudobombur Secretan, 1975
Type species: Pseudobombur nummuliticus Secretan,
1975.
Stratigraphic range: middle Eocene (Lutetian).
Pseudobombur nummuliticus Secretan, 1975
1975 - Pseudobombur nummuliticus Secretan; p. 332, Text-fig. 10,
PI. 11 (figs. 1-3), PI. 12 (fig. 2)
1 996 - Pseudobombur nummuliticus Secretan in Garassino & Teruzzi; p. 4
1999 - Pseudobombur nummuliticus Secretan in Beschin & Garassino;
p. 194
Holotype: MSNV 103.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported six specimens
(MSNV 102 bis, 103, 104, 106, 111 bis, 122) from Monte
Bolca (Verona).
Satyrocaris Garassino & Teruzzi, 1994
Type species: Satyrus cristatus Garassino &
Teruzzi, 1993.
Stratigraphic range: Upper Triassic (Norian).
Satyrocaris cristatus (Garassino & Teruzzi, 1993)
1993 - Satyrus cristatus Garassino & Teruzzi; p. 9, Text-figs. 12-14,
PI. 1 (fig. 4), PI. 2 (figs. 1-2)
1994 - Satyrocaris cristatus (Garassino & Teruzzi) in Garassino &
Teruzzi; p. 293
Holotype: MSNB 8190.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
8
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: Garassino & Teruzzi (1993) reported nine
specimens (MSNB 7790, 8190, 8240, 8248, 8299, 8338,
8341, 8350) from Ponte Giurino (Bergamo).
Indeterminates
Indeterminate penaeid
1993 - Indeterminate penaeid in Garassino & Teruzzi; p. 11, Text-
fig. 15, PI. 2 (fig. 3)
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported one
specimen (MSNB 7779) from Ponte Giurino (Bergamo).
Penaeid, genus and species indeterminate
1993 - Penaeid, genus and species indeterminate in Garassino &
Teruzzi; p. 12, Text-fig. 17
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported fìve
specimens (MSNB 7718, 7723, 7762, 7778; MSNM
il 0753) from Ponte Giurino (Bergamo).
Infraorder Caridea Dana, 1852
Family Oplophoridae Dana, 1852
Tonellocaris Garassino, 1 998
Type species: Tonellocaris brevirostrata Garassino,
1998.
Stratigraphic range: Lower Cretaceous (Barremian -
Aptian).
Tonellocaris brevirostrata Garassino, 1998
1998 - Tonellocaris brevirostrata Garassino; p. 64, Text-figs. 2-3
1999 - Tonellocaris brevirostrata Garassino in Bravi, Coppa, Garas¬
sino & Patricelli; p. 162
200 1 - Tonellocaris brevirostrata Garassino in Garassino & Teruzzi; p. 1 5 1
2002 - Tonellocaris brevirostrata Garassino in Garassino, Schram,
Taylor & Shen; p. 78
2003 - Tonellocaris brevirostrata Garassino in Garassino & Bravi; p. 590
2005 - Tonellocaris brevirostrata Garassino in Garassino & Jakobsen; p. 102
Holotype: MFSN 21537.
Stratigraphic range: Lower Cretaceous (Barremian -
Aptian).
Occurrence: Friuli-Venezia Giulia.
Material: Garassino (1998) reported three specimens
(MFSN 21537, 21540, 21541) from Torrente Comappo
(Udine).
Family Palaemonidae Rafinesque-Schmaltz, 1815
Alburnia Bravi & Garassino, 1998
Type species: Alburnia petinensis Bravi & Garassino,
1998.
Stratigraphic range: Lower Cretaceous (Albian).
Alburnia petinensis Bravi & Garassino, 1998
1998b - Alburnia petinensis Bravi & Garassino; p. 107, Text-
figs. 11-13
1998a - Alburnia petinensis Bravi & Garassino in Bravi & Garassino;
p. 155
1998 - Alburnia petinensis Bravi & Garassino in Garassino; p. 67
1999 - Alburnia petinensis Bravi & Garassino in Bravi, Coppa, Garas¬
sino & Patricelli; p. 162
2001 - Alburnia petinensis Bravi & Garassino in Garassino & Teruzzi;
p. 151
2002 - Alburnia petinensis Bravi & Garassino in Garassino, Schram,
Taylor & Shen; p. 78
2003 - Alburnia petinensis Bravi & Garassino in Garassino & Bravi;
p. 590
2005 - Alburnia petinensis Bravi & Garassino in Garassino & Jakob¬
sen; p. 102
Holotype: M 21834.
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998b) reported
three specimens (M 21834, 21835, 21836) from Petina
(Salerno).
Palaemon Weber, 1795
Type species: Palaemon adspersus Rathke, 1837.
Stratigraphic range: Lower Cretaceous (Aptian) -
Recent.
Palaemon antonellae Garassino & Bravi, 2003
2003 - Palaemon antonellae Garassino & Bravi; p. 589, Text-
figs. 1-2
2005 - Palaemon antonellae Garassino & Bravi in Garassino & Jakob¬
sen; p. 103
Holotype: PrC24.
Stratigraphic range: Lower Cretaceous (Aptian).
Occurrence: Campania.
Material: Garassino & Bravi (2003) reported 53
specimens (thè best specimens are: DGUN PrCl, PrC2,
PrC7, PrCl 3, PrC24) from Profeti (Caserta).
Palaemon fabricii Michelotti, 1861
1861 - Palaemon fabricii Michelotti; p. 141, PI. 14 (fig. 4)
1929 - Palaemon fabricii Michelotti in Glaessner; p. 291
Holotype: unknown.
Stratigraphic range: upper Oligocene (Chattian).
Occurrence: Veneto.
Material: Michelotti (1861) reported one specimen
from Salcedo (Vicenza) (repository and catalogue number
unknown).
Palaemon vesolensis Bravi, Coppa, Garassino &
Patricelli, 1999
1999 - Palaemon vesolensis Bravi, Coppa, Garassino & Patricelli;
p. 159, Text-figs. 13-17
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DEC APODA FROM 1TALY
9
2001 - Palaemon vesolensis Bravi, Coppa, Garassino & Patricelli in
Garassino & Teruzzi; p. 151
2002 - Palaemon vesolensis Bravi, Coppa, Garassino & Patricelli in
Garassino, Schram, Taylor & Shen; p. 78
2003 - Palaemon vesolensis Bravi, Coppa, Garassino & Patricelli in
Garassino & Bravi; p. 590
2005 - Palaemon vesolensis Bravi, Coppa, Garassino & Patricelli in
Garassino & Jakobsen; p. 103
Holotype: A 4491/8.
Stratigraphic range: Upper Cretaceous (Campanian-
Maastrichtian).
Occurrence: Campania.
Material: Bravi et al. (1999) reported 93 specimens
(A 4491/from 1 to 31, 34, 41; MSNM from Ì24731 to
Ì24796) from Monte Vesole (Salerno).
Palaemon sp.
1892c - Palaemon sp. in Ristori; p. 160
1 9 1 Oa - Palaemon sp. in Fabiani; p. 18
Stratigraphic range: upper Oligocene (Rupelian).
Occurrence: Veneto.
Material: Ristori (1892c) and Fabiani (1910a)
reported some specimens from Chiavon (Vicenza)
(repository and catalogue number unknown).
Indeterminate family
Acanthinopus Pinna, 1974
Type species: Acanthinopus gibbosus Pinna, 1974.
Stratigraphic range: Upper Triassic (Norian).
Acanthinopus gibbosus Pinna, 1 974
1974 - Acanthinopus gibbosus Pinna; p. 23, PI. 12 (fig. 3), PI. 13
(fig- 5)
1996 - Acanthinopus gibbosus Pinna in Garassino, Teruzzi & Dalla
Vecchia; p. 32
1996 - Acanthinopus gibbosus Pinna in Garassino & Teruzzi; p. 8
1997 - Acanthinopus gibbosus Pinna in Garassino; p. 107
1998a - Acanthinopus gibbosus Pinna in Bravi & Garassino;
p. 155
1 99 8b - Acanthinopus gibbosus Pinna in Bravi & Garassino;
p. 108
1998 - Acanthinopus gibbosus Pinna in Garassino; p. 65
1999 - Acanthinopus gibbosus Pinna in Bravi, Coppa, Garassino &
Patricelli; p. 161
2001 - Acanthinopus gibbosus Pinna in Garassino & Teruzzi; p. 150
2002 - Acanthinopus gibbosus Pinna in Garassino, Schram, Taylor &
Shen; p. 76
2003 - Acanthinopus gibbosus Pinna in Garassino & Bravi; p. 590
2005 - Acanthinopus gibbosus Pinna in Garassino & Jakobsen; p. 102
Holotype: MSNB3109.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia, Friuli-Venezia Giulia.
Material: Pinna (1974) reported one specimen
(MSNB 3109) from Cene (Bergamo); Garassino et al.
(1996) reported two specimens (MFSN 16104 a-b, 18454
a-b) from Rio Seazza (Udine).
Leiothorax Pinna, 1974
Type species: Leiothorax triasicus Pinna, 1974.
Stratigraphic range: Upper Triassic (Norian).
Leiothorax triasicus Pinna, 1974
1974 - Leiothorax triasicus Pinna; p. 24, PI. 12 (fig. 2), PI. 13 (fig. 4)
1996 - Leiothorax triasicus Pinna in Garassino, Teruzzi & Dalla Vec¬
chia; p. 33
1996 - Leiothorax triasicus Pinna in Garassino & Teruzzi; p. 8
1997 - Leiothorax triasicus Pinna in Garassino; p. 107
1998a - Leiothorax triasicus Pinna in Bravi & Garassino; p. 155
1 998b - Leiothorax triasicus Pinna in Bravi & Garassino; p. 108
1998 - Leiothorax triasicus Pinna in Garassino; p. 65
1999 - Leiothorax triasicus Pinna in Bravi, Coppa, Garassino & Patri¬
celli; p. 161
2001 - Leiothorax triasicus Pinna in Garassino & Teruzzi; p. 150
2002 - Leiothorax triasicus Pinna in Garassino, Schram, Taylor &
Shen; p. 76
2003 - Leiothorax triasicus Pinna in Garassino & Bravi; p. 590
2005 - Leiothorax triasicus Pinna in Garassino & Jakobsen; p. 102
Holotype: MSNB 3156.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Pinna (1974) reported one specimen
(MSNB 3156) from Cene (Bergamo).
Parvocaris Bravi & Garassino, 1 998
Type species: Parvocaris samnitica Bravi &
Garassino, 1998.
Stratigraphic range: Lower Cretaceous (Albian).
Parvocaris samnitica Bravi & Garassino, 1 998
1998a - Parvocaris samnitica Bravi & Garassino; p. 1 54, Text-figs. 24-25
1998b - Parvocaris samnitica Bravi & Garassino in Bravi & Garas¬
sino; p. 109
1998 - Parvocaris samnitica Bravi & Garassino in Garassino; p. 67
1999 - Parvocaris samnitica Bravi & Garassino in Bravi, Coppa,
Garassino & Patricelli; p. 162
2001 - Parvocaris samnitica Bravi & Garassino in Garassino &
Teruzzi; p. 151
2002 - Parvocaris samnitica Bravi & Garassino in Garassino, Schram,
Taylor & Shen; p. 76
2003 - Parvocaris samnitica Bravi & Garassino in Garassino & Bravi;
p. 590
2005 - Parvocaris samnitica Bravi & Garassino in Garassino & Jakob¬
sen; p. 102
Holotype: M 20545.
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998a) reported 14
specimens (M 19309, 19334, 19344, 19373, 19389,
20525, 20545, 20571, 20723, 20772, 20773, St5/29, St7/
7, St7/8) from Pietraroia (Benevento).
Pinnacaris Garassino & Teruzzi, 1993
Type species: Pinnacaris dentata Garassino &
Teruzzi, 1993.
10
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: Upper Triassic (Norian).
Pinnacaris dentata Garassino & Teruzzi, 1993
1993 - Pinnacaris dentata Garassino & Teruzzi; p. 12, Text-figs. 1 8-
19, PI. 2(fig. 4)
1996 - Pinnacaris dentata Garassino & Teruzzi in Garassino &
Teruzzi; p. 8
1997 - Pinnacaris dentata Garassino & Teruzzi in Garassino; p. 107
1 998a - Pinnacaris dentata Garassino & Teruzzi in Bravi & Garassino;
p. 155
1 998b -Pinnacaris dentata Garassino & Teruzzi in Bravi & Garassino;
p. 108
1998 - Pinnacaris dentata Garassino & Teruzzi in Garassino; p. 65
1999 - Pinnacaris dentata Garassino & Teruzzi in Bravi, Coppa,
Garassino & Patricelli; p. 161
2001 - Pinnacaris dentata Garassino & Teruzzi in Garassino &
Teruzzi; p. 150
2002 - Pinnacaris dentata Garassino & Teruzzi in Garassino, Schram,
Taylor & Shen; p. 76
2003 - Pinnacaris dentata Garassino & Teruzzi in Garassino & Bravi;
p. 590
2005 - Pinnacaris dentata Garassino & Teruzzi in Garassino & Jakob-
sen; p. 102
Holotype: MSNM il 0691 -il 0692 (part and counter-
part).
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported four
specimens (MSNB 7693, 7731; MSNM il 0691 -il 0692,
il 0693) from Ponte Giurino (Bergamo).
Indeterminates
Indeterminate caridean
1995 - Indeterminate caridean in Garassino & Teruzzi; p. 83, Text-fig. 3
Stratigraphic range: Lower Cretaceous (Hauterivi-
an-Barremian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino & Teruzzi (1995) reported two
specimens (MFSN 4309, 4316) from Vemasso (Udine).
Caridean genus et species indeterminate
1 996 - Caridean gen. et sp. indet. in Garassino, Teruzzi & Dalla Vec¬
chia; p. 32, Text-fig. 16
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino et al. (1996) reported one speci¬
men (MFSN 16140) from Caprizzi (Udine).
Infraorder Astacidea Latreille, 1802
Family Glypheidae Zittel, 1885
Glyphea v. Meyer, 1835
Type species: Palinurus regleyanus Desmarest, 1822.
Stratigraphic range: Upper Triassic (Camian) -
Eocene (Bartonian).
Glyphea rigoi Garassino, 2000
1996 - Glyphea sp. in Garassino, Teruzzi & Dalla Vecchia; p. 43
2000a - Glyphea rigoi Garassino; p. 61, Text-fig. 1
Holotype: MFSN 22976.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Friuli-Venezia Giulia.
Material: Garassino et al. (1996) reported five speci¬
mens (MFSN 1414 a-b, 1953, 1954, 16095 a-b, 16182)
from Caprizzi and Rio Seazza (Udine); Garassino (2000a)
reported one specimen (MFSN 22976) from Caprizzi
(Udine).
Glyphea tonelloi Garassino, 1997
1997 - Glyphea tonelloi Garassino; p. 87, Text-figs. 2-5
Holotype: MFSN 19863.
Stratigraphic range: Lower Cretaceous (Barremian -
Aptian).
Occurrence: Friuli-Venzia Giulia.
Material: Garassino (1997) reported four specimens
(MFSN 19859, 19861, 19862, 19863) from Torrente Cor-
nappo (Udine).
Glyphea Incannata Garassino, 1 996
1996 - Glyphea tricarinata Garassino; p. 341, Text-figs. 6-8, 22-25
Holotype: MSNM Ì7609.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino (1996) reported 137 specimens
(thè best specimens are: MSNM Ì7609, Ì9938, Ì9942,
Ì9944, Ì9946, Ì9963, Ì9967, Ì9970, Ì9980, Ì9997, il 0007,
il 0451, il 0455) from Osteno (Como).
fFamily Mecochiridae Van Straelen, 1925
Mecochirus Germar, 1827
Type species: Macrourites longimanatus Schlotheim,
1820.
Stratigraphic range: Lower Jurassic (Sinemurian) -
Upper Cretaceous (Maastrichtian).
Mecochirus germari Garassino, 1 996
1996 - Mecochirus germari Garassino; p. 346, Text-figs. 9-10,
26-29
Holotype: MSNM il 3521 a-b.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino (1996) reported 81 specimens
(thè best specimens are: MSNM il 01 19, i 1 0 1 2 1 , il 01 24,
i 1 2 1 32 a-b, Ì10136, Ì10139, Ì10141, Ì10172, Ì10294,
il 0920, il 3520, il 3521 a-b) from Osteno (Como).
Pseudoglyphea Oppel, 1861
Type species: Glyphea grandis v. Meyer, 1837.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
11
Stratigraphic range: Upper Triassic (Camian) -
Upper Jurassic (Oxfordian).
Pseudoglyphea ancylochelis (Woodward, 1863)
1863 - Scapiteli s ancylochelis Woodward; p. 3 1 9, Text-fig. 1 1
1924 - Scapheus ancylochelis Woodward in Van Straelen; p. 224,
Text-fig. 3
1 925 - Scapheus ancylochelis Woodward in Van Straelen; p. 2 1 0, Text-
fig. 104, PI. 7(fig. 2)
1925 - Pseudoglyphea ancylochelis (Woodward) in Woods; p. 46,
PI. 12 (figs. 2-3)
1929 - Pseudoglyphea ancylochelis (Woodward) in Glaessner; p. 353
1996 - Pseudoglyphea ancylochelis (Woodward) in Garassino; p. 349,
Text-figs. 11-14, 30-31
Holotype: NHM 46322.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino (1996) reported seven specimens
(MSNM Ì10149, i 1 0 1 69, Ì10254, Ì10256-Ì10293, Ì10258,
il 0264, il 0295) from Osteno (Como).
Note: this species is also reported from thè Lower
Jurassic (Sinemurian) of Lyme Regis (Great Britain).
Pseudoglyphea gigantea Garassino & Teruzzi, 1993
1993 - Pseudoglyphea gigantea Garassino & Teruzzi; p. 14, Text-
figs. 20-28, PI. 3 (figs. 1-4)
Holotype: MSNM il 0678-i 10679 (part and counter-
part).
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported 183
specimens (thè best specimens are: MSNB 7648, 7561,
7558-7554, 7624, 7659-7661, 7573, 7620-7621, 7562-
7702, 7610, 7684, 8252, 8336, 8241, 8348, 8321; MSNM
from il 0676 to il 0681, il 0689-i 10690) from Ponte
Giurino (Bergamo).
Pseudoglyphea paronae (Colosi, 1921)
1921 - Heteroglyphea paronae Colosi; p. 81, Text-figs. 1-2
1 924 - Pseudoglyphea paronae (Colosi) in Van Straelen; p. 226, Text-
fig. 6
1 925 - Pseudoglyphea paronae (Colosi) in Van Straelen; p. 205
1929 - Pseudoglyphea paronai (Colosi) in Glaessner; p. 356
1969 - Pseudoglyphea paronae (Colosi) in Glaessner; p. 466
1996 - Pseudoglyphea paronae (Colosi) in Garassino; p. 353
Holotype: lost.
Stratigraphic range: Jurassic.
Occurrence: Liguria.
Material: Colosi (1921) reported one specimen from
La Spezia, today lost.
fFamily Erymidae Van Straelen, 1924
Eryma v. Meyer, 1 840
Type species: Macrourites modestiformis Schlotheim,
1822.
Stratigraphic range: Lower Jurassic (Sinemurian) -
Upper Cretaceous (Cenomanian).
Eryma meyeri Garassino, 1996
1996 -Eryma meyeri Garassino; p. 335, Text-figs. 1-3, 15-17
Holotype: MSNM Ì7606.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Garassino (1996) reported 41 specimens
(thè best specimens are: MSNM Ì7606, Ì7607, Ì9871,
Ì9875, Ì9884, Ì9886, Ì9893, Ì9895, Ì9896, Ì9902, Ì9915,
i99 1 6, il 0265, il 0635, il 0656) from Osteno (Como).
? Eryma rinellincola De Gregorio, 1884
1884 - ? Eryma rinellincola De Gregorio; p. 134
1929 - Eryma rinellincola De Gregorio in Glaessner; p. 159
Holotype: lost.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: De Gregorio (1884) reported one specimen
from Contrada Rotoli (Palermo), today lost.
Pustulina Quenstedt, 1857
Type species: Pustulina suevica Quenstedt, 1857.
Stratigraphic range: Lower Jurassic (Sinemurian) -
Upper Cretaceous (Campanian - Maastrichtian).
Pustulina sinemuriana (Garassino, 1996)
1996 - Phlyctisoma sinemuriana Garassino; p. 338, Text-figs. 4-5,
18-21
Holotype: MSNMÌ13517.
Stratigraphic range: Lower Jurassic (Sinemurinan).
Occurrence: Lombardia.
Material: Garassino (1996) reported ten specimens
(thè best specimens are: MSNM Ì7608-Ì99 1 1 , Ì9907-
Ì9887, Ì9909-Ì9873, il 0450, il 35 17) from Osteno
(Como).
Family Nephropidae Dana, 1852
Hoploparia McCoy, 1 849
Type species: Astacus longimanus Sowerby, 1826.
Stratigraphic range: Lower Cretaceous (Hauterivi-
an) - Miocene.
Hoploparia sp.
1889 - Hoploparia sp. in Ristori; p. 41 1, PI. 1 5 (fig. 19)
1929 - Hoploparia sp. in Glaessner; p. 223
Stratigraphic range: Miocene.
Occurrence: Liguria.
Material: Ristori (1889) reported two specimens from
Mioglia and S. Giustina (Savona), housed in don Perran-
do’s collection, today lost.
12
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Hoploparia sp.
2005 - Hoploparia sp. in Beschin, De Angeli, Checchi & Zarantonello;
p. 7, Text-fig. 2, PI. 1 (fig. 1 a-c)
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported one speci¬
men (MCZ 2360) from Grola di Comedo Vicentino
(Vicenza).
Metanephrops Jenkins, 1972
Type species: Nephrops japonicus Tapparone-Cane-
fri, 1873.
Stratigraphic range: Upper Cretaceous (Maastrich-
tian) - Recent.
Metanephrops sp.
2003 - Metanephrops sp. in Garassino, De Angeli & De Polli; p. 387,
Text-figs. 4-5
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Garassino et al. (2003) reported one speci¬
men (MSNM Ì26261) from “Fontanella” di Grancona
(Vicenza).
Nephropsis Wood-Mason, 1873
Type species: Nephropsis stewarti Wood-Mason, 1873.
Stratigraphic range: Eocene - Recent.
Nephropsis sp.
2004a - Nephropsis sp. in Garassino & De Angeli; p. 35, Text-fig. 2
Stratigraphic range: Pleistocene.
Occurrence: Emilia Romagna.
Material: Garassino & De Angeli (2004a) reported
one specimen (MG 0620) from Fiume Enza (Parma).
fFamily Platychelidae Glaessner, 1969
Glaessnericaris Garassino & Teruzzi, 1993
Type species: Glaessnericaris macrochela Garassino
& Teruzzi, 1993.
Stratigraphic range: Upper Triassic (Norian).
Glaessnericaris dubia (Pinna, 1974)
1974 - Protoclytiopsis dubia Pinna; p. 26, PI. 1 (figs. 1-3), PI. 16
(figs. 1-3)
1993 - Glaessnericaris dubia (Pinna) in Garassino & Teruzzi; p. 24
Holotype: MSNB 3186.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Pinna (1974) reported nine specimens
(MSNB 3186, 3187, 3188, 3155, 3183, 3194; MSNM
Ì4475, Ì4479) from Cene (Bergamo).
Glaessnericaris macrochela Garassino & Teruzzi,
1993
1993 - Glaessnericaris macrochela Garassino & Teruzzi; p. 2 1 , Text-
figs. 41-48, PI. 5 (figs. 1-4)
Holotype: MSNB 4202.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (1993) reported
310 specimens (thè best specimens are: MSNB 4202,
4205, 4208, 7564, 7556-7553, 7591, 7701, 7583-7627,
7622, 7679, 7642, 7640, 7652, 7672, 7732, 7557, 7572,
7559 a-b, 7708-7710, 8214, 8315, 8249, 8237; MSNM
i 1 0732, i 1 0727, i 1 0736, i 1 073 1 ) from Ponte Giurino
(Bergamo).
Glaessnericaris sp.
1996 - Glaessnericaris sp. in Garassino, Teruzzi & Dalla Vecchia;
p. 44
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Friuli- Venzia Giulia.
Material: Garassino et al. (1996) reported one speci¬
men (MFSN 16146 a-b) from Caprizzi (Udine).
fFamily Uncinidae Beurlen, 1928
Uncina Quenstedt, 1 85 1
Type species: Uncina posidoniae Quenstedt, 1851.
Stratigraphic range: Lower Jurassic (Sinemurian/
Pliensbachian - Toarcian).
Uncina alpina Schweigert, Garassino, Hall, Hauff &
Karasawa, 2003
2001 - Uncina cfr. U. posidoniae Quenstedt in Garassino & Teruzzi;
p. 189, Text-fig. 2
2003 - Uncina alpina Schweigert, Garassino, Hall, Hauff & Kara¬
sawa; p. 9, Text-fig. 5 a, PI. 10 (figs. 1-5)
Holotype: MSNM il 0864.
Stratigraphic range: Lower Jurassic (Toarcian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (2001) reported three
specimens (MSNM il 0851, il 0863, il 0864) from Sogno
(Bergamo); Schweigert et al. (2003) described this spe¬
cies on thè three specimens previously studied by Garas¬
sino & Teruzzi.
Indeterminates
Family, genus et species indeterminate
1996 - Family, gen. et sp. indet. in Garassino, Teruzzi & Dalla Vecchia;
p. 44, Text-fig. 23
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino et al. (1996) reported one speci¬
men (MFSN 1413) from Caprizzi (Udine).
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
13
Family, genus et species indeterminate
1998a - Family, gen. et sp. indet. in Bravi & Garassino; p. 156
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998a) reported two
specimens (M 20547, 20724) fforn Pietraroia (Benevento).
Family ?Platichelidae, genus et species indetenninate
1996 - Family ?Platichelidae, gen. et sp. indet. in Garassino, Teruzzi &
Dalla Vecchia; p. 45, Text-fig. 24
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino et al. (1996) reported one speci¬
men (MFSN 16170 a-b) from Monte Auda (Udine).
Genus et species indeterminate
1999 - Gen. et sp. indet. in Garassino; p. 63, Text-fig. 1
Stratigraphic range: Lower Cretaceous (Hauterivian-
Barremian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino (1999) reported two specimens
(MFSN 22977, 22978 a-b) from Vemasso (Udine).
Infraorder Thalassinidea Latreille, 1 83 1
Superfamily Thalassinoidea Latreille, 1831
Family Thalassinidae Latreille, 1831
Thalassina Latreille, 1806
Type species: Thalassina scorpionides Latreille, 1806.
Stratigraphic range: Pleistocene - Recent.
Thalassina sp.
1 89 la— Thalassina sp. in Ristori; p. 14, PI. 1 (figs. 16-17)
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (189 la) reported two specimens
from Spicchio (Empoli), today lost.
Superfamily Callianassoidea Dana, 1852
Family Callianassidae Dana, 1852
Callianassa Leach, 1814
Type species: Cancer ( Astacus ) subterraneus
Montagu, 1808.
Stratigraphic range: Upper Cretaceous - Recent.
Callianassa calaritana Ristori, 1896
1896 - Callianassa calaritana Ristori; p. 512, PI. 12 (fig. 9)
1902 - Callianassa calaritana Ristori in Lovisato; p. 8
1909 - Callianassa calaritana Ristori in Lòrenthey; p. 218
1929 - Callianassa calaritana Ristori in Glaessner; p. 77
1950 - Callianassa calaritana Ristori in Comaschi Caria; p. 326
1956 - Callianassa calaritana Ristori in Comaschi Caria; p. 288
Holotype: lost.
Stratigraphic range: Pleistocene.
Occurrence: Sardegna.
Material: Ristori (1896) reported one specimen
from S. Avendrace (Cagliari), today lost; Lovisato
(1902) reported some specimens from S. Bartolomeo
(Cagliari) (repository and catalogue number unknown);
Lòrenthey (1909) reported one specimen from S. Aven¬
drace (Cagliari) (repository and catalogue number
unknown).
Callianassa canavarii Ristori, 1889
1889 - Callianassa canavarii Ristori; p. 409, PI. 15 (figs. 17-18)
1929 - Callianassa canavarii Ristori in Glaessner; p. 77
1974 - Callianassa canavarii Ristori in Mastrorilli; p. 4
Holotype: lost.
Stratigraphic range: Oligocene.
Occurrence: Liguria.
Material: Ristori (1889) reported two specimens from
S. Giustina and Sassello (Savona), housed in don Perran-
do’s collection, today lost.
Callianassa desmarestiana A. Milne Edwards, 1860
1829 - Pagurus desmarestianus de Serres ( nomen nudum); p. 154
1860 - Callianassa desmarestiana A. Milne Edwards; p. 335, PI. 13
(fig- 4)
1888 - Callianassa desmarestiana (A. Milne Edwards) in Ristori; p.
217, PI. 4 (figs. 12-13)
1 896 - Callianassa desmarestiana (A. Milne Edwards) in Ristori; p. 5 1 3
1 902 - Callianassa desmarestiana (A. Milne Edwards) in Lovisato; p. 7
1909 - Callianassa desmarestiana (A. Milne Edwards) in Lòrenthey;
p. 213, PI. 1 (figs. 3-4)
1 929 -Callianassa desmarestiana(A. Milne Edwards) in Glaessner; p. 79
1950 - Callianassa desmarestiana (A. Milne Edwards) in Comaschi
Caria; p. 326
1956 - Callianassa desmarestiana (A. Milne Edwards) in Comaschi
Caria; p. 288
Holotype: unknown.
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Ristori (1888) reported some specimens
from S. Bartolomeo (Cagliari), housed in Lovisato’s
collection (repository and catalogue number unknown);
Ristori (1896) reported one specimen from Bosa (Nuoro)
(repository and catalogue number unknown); Lovisato
(1902) reported some specimens from S. Bartolomeo
(Cagliari) (repository and catalogue number unknown);
Lòrenthey (1909) reported one specimen from S. Bar¬
tolomeo (Cagliari), housed in Lovisato’s collection
(repository and catalogue number unknown).
Note: this species is also reported from thè Miocene
of France.
Callianassa cfr. C.ferox Bittner, 1893
2005 - Callianassa cfr. C.ferox Bittner in Beschin, De Angeli, Checchi
& Zarantonello; p. 8, PI. 1 (figs. 2-3)
14
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported two speci-
mens (MCZ 2371, 2372) from Grola di Comedo Vicen¬
tino (Vicenza).
Callianassa michelotti A. Milne Edwards, 1860
1860 - Callianassa michelotti A. Milne Edwards; p. 341, PI. 14
(fig- 3)
1871 - Callianassa michelotti A. Milne Edwards in Fritsch; p. 691, PI.
17 (figs. 5-13)
1886 - Callianassa michelotti A. Milne Edwards in Noetling; p. 84,
PI. 5 (fig. 4)
1895 - Callianassa michelotti A. Milne Edwards in Crema; p. 667,
Text-fig. 3
1903 - Callianassa michelotti A. Milne Edwards in Kinkelin; p. 31
1928 - Callianassa michelotti A. Milne Edwards in Glaessner; p. 167
1929 - Callianassa michelotti A. Milne Edwards in Glaessner; p. 85
Holotype: MNHN B32690.
Stratigraphic range: lower Oligocene (Rupelian) -
Miocene.
Occurrence: Piemonte.
Material: A. Milne Edwards (1860) reported one
specimen (MNHN B32690) from Superga (Torino);
Crema (1895) reported some specimens from Torino
(repository and catalogue number unknown).
Note: this species is also reported from thè Oligocene
of Germany and thè Miocene of Austria.
Callianassa pedemontana Crema, 1895
1895 - Callianassa pedemontana Crema; p. 665, Text-fig. 1 a-b
1 909 - Callianassa pedemontana Crema in Lòrenthey; p. 220
1 929 - Callianassa pedemontana Crema in Glaessner; p. 87
Holotype: lost.
Stratigraphic range: Miocene.
Occurrence: Piemonte, Sardegna.
Material: Crema (1895) reported one specimen from
Torino, today lost; Lòrenthey (1909) reported one speci¬
men from Coroneddu (Cagliari), housed in Lovisato’s
collection (repository and catalogue number unknown).
Callianassa cfr. C. rakosiensis Lòrenthey, 1 898
1909 - Callianassa cfr. C. rakosiensis Lòrenthey in Lòrenthey; p. 217
1 929 - Callianassa cfr. C. rakosiensis Lòrenthey in Glaessner; p. 89
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported some specimens
from Torralba (Sassari) (repository and catalogue number
unknown).
Callianassa rovasendae Crema, 1 895
1895 - Callianassa rovasendae Crema; p. 666, Text-fig. 2 a-b
1929 - Callianassa rovasendae Crema in Glaessner; p. 89
Holotype: lost.
Stratigraphic range: Miocene.
Occurrence: Piemonte.
Material: Crema (1895) reported one specimen from
Baldissero (Torino), today lost.
Callianassa sismondai A. Milne Edwards, 1 860
1 846 - Grapsus sp. in E. Sismonda; p. 69, PI. 3 (fig. 7)
1860 - Callianassa sismondai A. Milne Edwards; p. 342, PI. 14
(fig- 4)
1861 - Callianassa sismondae A. Milne Edwards in E. Sismonda;
p. 21
1895 - Callianassa sismondae A. Milne Edwards in Crema; p. 667,
Text-figs. 4-6, 7 a-b
1928 - Callianassa sismondae A. Milne Edwards in Glaessner; p. 168
1929 - Callianassa sismondae A. Milne Edwards in Glaessner; p. 90
Holotype: MNHN B32689.
Stratigraphic range: Miocene.
Occurrence: Piemonte.
Material: A. Milne Edwards (1860) reported one
specimen (MNHN B32689) from Torino; E. Sismonda
(1861) reported some specimens from Torino (repository
and catalogue number unknown); Crema (1895) reported
some specimens from Torino (repository and catalogue
number unknown).
Note: E. Sismonda (1846) reported one specimen
from Torino, ascribing it to Grapsus and later E. Sis¬
monda (1861) ascribed thè same specimen to Callianassa
sismondai. This species is also reported from thè Miocene
of Austria.
Callianassa subterranea (Montagu, 1808)
1808 - Callianassa subterranea Montagu; p. 88, PI. 3 (figs. 1-2)
1 895 - Callianassa subterranea Montagu in Crema; p. 669, Text-
fig. 8 a-b
1909 - Callianassa subterranea Montagu in Lòrenthey; p. 219, PI. 1
(fig- 11)
1929 - Callinassa subterranea Montagu in Glaessner; p. 91
Holotype: unknown.
Stratigraphic range: Miocene.
Occurrence: Piemonte, Sardegna.
Material: Crema (1895) reported one specimen from
Albugnano (Asti) (repository and catalogue number
unknown); Lòrenthey (1909) reported one specimen
from Alghero (Sassari) (repository and catalogue number
unknown).
Callianassa subterranea var. dentata Ristori, 1891
1891 b - Callianassa subterranea var. dentata Ristori; p. 24, PI. 1
(figs. 19-20)
1 895 - Callianassa subterranea var. dentata Ristori in Crema; p. 670
1 9 1 4 - Callianassa subterranea var. dentata Ristori in M. Gemmellaro;
p. 94, PI. 1 (figs. 31-32)
1929 - Callianassa subterranea var. dentata Ristori in Glaessner;
p. 91
1981 - Callianassa subterranea var. dentata Ristori in Varola; p. 10,
PI. 1 (figs. 1-2,4), PI. 2 (figs. 1,3-4)
Stratigraphic range: Miocene - Pleistocene (Sici-
lian).
Occurrence: Piemonte, Lazio, Puglia, Sicilia.
Material: Ristori ( 1 89 1 b) reported one specimen from
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
15
Astura (Roma), housed in Clerici’s collection (repository
and catalogue nurnber unknown); Crema (1895) reported
some specimens from Torino (repository and catalogue
nurnber unknown); M. Gemmellaro (1914) reported two
specimens from Monte Pellegrino (MGUP, catalogue
nurnber unknown); Varola (1981) reported one specimen
from Porto Craulo (Otranto) (GNSL, catalogue nurnber
unknown).
Callianassa sp.
1 889 - Callianassa sp. in Ristori; p. 410
1929 - Callianassa sp. in Glaessner; p. 96
1974 - Callianassa sp. in Mastrorilli; p. 4
Stratigraphic range: Miocene.
Occurrence: Liguria.
Material: Ristori (1889) reported many specimens
from S. Giustina and Sassello (Savona), today lost.
Callianassa sp.
1 895 - Callianassa sp. in Crema; p. 670, Text-figs. 9 a-b, 1 0 a-b
1929 - Callianassa sp. in Glaessner; p. 96
Stratigraphic range: Miocene.
Occurrence: Piemonte.
Material: Crema (1895) reported some specimens
from Torino, Sciolze and Baldissero (Torino), today lost.
Callianassa sp.
1909 - Callianassa sp. in Lòrenthey; p. 222
1929 - Callianassa sp. in Glaessner; p. 96
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material Lòrenthey (1909) reported some specimens
from S. Lorenzo di Nulvi (Sassari) (repository and cata¬
logue nurnber unknown).
Callianassa sp.
1991 - Callianassa sp. in Marras & Ventura; p. 107, PI. 2 (fig. 3)
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Marras & Ventura (1991) reported one
specimen from Sassari (repository and catalogue nurnber
unknown).
Corallianassa Manning, 1987
Type species: Callianassa longiventris A. Milne
Edwards, 1870.
Stratigraphic range: Upper Cretaceous - Recent.
Corallianassa rigoi De Angeli & Garassino, 2006
2006a - Corallianassa rigoi De Angeli & Garassino; p. 270, Text-
figs. 2, 3 (a-c)
Holotype: MSNM Ì26581.
Stratigraphic range: Cretaceous (Aptian-Campanian).
Occurrence: Friuli-Venezia Giulia.
Material: De Angeli & Garassino (2006a) reported
six specimens (MSNM from Ì26578 to Ì26583) from
Monte Ciaurlec (Pordenone).
Eoglypturus Beschin, De Angeli, Checchi &
Zarantonello, 2005
Type species: Eoglypturus grolensis Beschin,
De Angeli, Checchi & Zarantonello, 2005.
Stratigraphic range: middle Eocene (Lutetian).
Eoglypturus grolensis Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 - Eoglypturus grolensis Beschin, De Angeli, Checchi & Zaran¬
tonello; p. 10, Text-fig. 5, PI. 1 (fig. 6 a-b)
Holotype: MCZ 2381 (I.G. 296600).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported one speci¬
men (MCZ 2381) from Grola di Comedo Vicentino
(Vicenza).
Eucalliax Manning & Felder, 1991
Type species: Callianassa quadracuta Biffar, 1970.
Stratigraphic range: Eocene - Recent.
Eucalliax vicetina Beschin, Busulini, De Angeli &
Tessier, 2002
2002 - Eucalliax vicetina Beschin, Busulini, De Angeli & Tessier;
p. 10, Text-fig. 4, PI. 1 (figs. 1-3)
2004 - Eucalliax vicetina Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 13
2005 - Eucalliax vicetina Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli, Checchi & Zarantonello; p. 9, Text-fig. 4,
PI. 1 (fig- 5)
Holotype: MCZ 2259 (I.G. 296387).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported five speci¬
mens (MCZ 1187, 2259, 2260, 2261, 2262) from “Main”
quarry of Arzignano (Vicenza); Beschin et al. (2005)
reported one specimen (MCZ 2363) from Grola di
Comedo Vicentino (Vicenza).
Neocallichirus Sakai, 1988
Type species: Neocallichirus horneri Sakai, 1988.
Stratigraphic range: Eocene - Recent.
Neocallichirus allegranzii Beschin, De Angeli, Checchi
& Zarantonello, 2005
2005 - Neocallichirus allegranzii Beschin, De Angeli, Checchi &
Zarantonello; p. 8, Text-fig. 3, PI. 1 (figs. 7-8)
Holotype: MCZ 2318 (I.G. 296537).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
16
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: Beschin et al. (2005) reported seven speci¬
men (MCZ 2281, 2282, 2306, 2318, 2321, 2329, 2385)
from Grola di Comedo Vicentino (Vicenza).
Neocallichirus borensis Beschin, De Angeli, Checchi &
Mietto, 2006
2006 - Neocallichirus borensis Beschin, De Angeli, Checchi &
Mietto; p. 97, Text-fig. 2, PI. (figs. 4 a-b, 5-6)
Holotype: MCZ 2423.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (2006) reported ten speci¬
men (MCZ from 2423 to 2428, from 2450 to 2453) from
Mulino del Boro (Vicenza).
Neocallichirus fortisi Beschin, Busulini, De Angeli &
Tessier, 2002
2002 - Neocallichirus fortisi Beschin, Busulini, De Angeli & Tessier;
p. 9, Text-fig. 3, PI. 1 (fig. 4 a-c)
2004 - Neocallichirus fortisi Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 113
2005 - Neocallichirus fortisi Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli, Checchi & Zarantonello; p. 8, PI. 1 (fig. 4)
Holotype: MCZ 2258 (I.G. 296386).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported two speci¬
men (MCZ 2258 a-b) from “Main” quarry of Arzignano
(Vicenza); Beschin et al. (2005) reported one specimen
(MCZ 2328) from Grola di Comedo Vicentino (Vicen¬
za).
Neocallichirus sp.
1995 - Callianassa sp. in De Angeli; p. 10, Text-fig. 2 (1), PI. 1 (figs. 1-2)
2001 - Callianassa sp. in De Angeli & Beschin; p. 10
2006 - Neocallichirus sp. in Beschin, De Angeli, Checchi & Mietto;
p. 98, Text-fig. 3, PI. 1 (figs. 1-3)
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli (1995) reported four specimens
(MCZ from 1486 to 1489) from “Fontanella” di Grancona
(Vicenza); Beschin et al. (2006) reported three specimens
(MCZ 2420, 2421, 2422) from Buso della Rana (Vicenza).
Family Ctenochelidae Manning & Felder, 1991
Callianopsis de Saint Laurent, 1973
Type species: Callianassa goniophthalma Rathbun,
1902.
Stratigraphic range: Eocene - Recent.
Callianopsis microspineus Beschin, De Angeli, Checchi
& Zarantonello, 2005
2005 - Callianopsis microspineus Beschin, De Angeli, Checchi &
Zarantonello; p. 1 1, Text-fig. 7, PI. 2 (fig. 1 a-b)
Holotype: MCZ 2373 (I.G. 296592).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported two speci¬
mens (MCZ 2323, 2373) from Grola di Comedo Vicen¬
tino (Vicenza).
Ctenocheles Kishinouye, 1926
Type species: Ctenocheles balssi Kishinouye, 1926.
Stratigraphic range: Upper Cretaceous - Recent.
Ctenocheles ornatus Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 - Ctenocheles ornatus Beschin, De Angeli, Checchi & Zaran¬
tonello; p. 10, Text-fig. 6, PI. 1 (figs. 9-11)
Holotype: MCZ 2334 (I.G. 296553).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported five speci¬
mens (MCZ 2283, 2309, 2310, 2332, 2334) from Grola di
Comedo Vicentino (Vicenza).
Ctenocheles valdellae (Fabiani, 1908)
1905 - Ilia sp. in Fabiani; p. 32
1908 - Ilia? Valdellae Fabiani; p. 21 1, 236, PI. 6 (fig. 15)
191 Oa — Ilia Valdellae Fabiani in Fabiani; p. 21, 33
1915 - Ilia Valdellae Fabiani in Fabiani; p. 284, 285
1929 - Ilia valdellae Fabiani in Glaessner; p. 225
1998 - Ctenocheles valdellae (Fabiani) in Beschin, Busulini,
De Angeli, Tessier & Ungaro; p. 15
2001 - Ctenocheles valdellae (Fabiani) in De Angeli & Beschin;
p. 10
2005 - Ctenocheles valdellae (Fabiani) in Beschin, De Angeli, Checchi
& Zarantonello; p. 1 1
Holotype: MGPD 23453.
Stratigraphic range: lower/middle Eocene, lower
Oligocene.
Occurrence: Veneto.
Material: Fabiani (1908, 191 Oa) reported two speci¬
mens (MGPD 23453, 23454) from Colle Valdella di
Nanto, Montruglio, and Monte Berico (Vicenza).
Ctenocheles sp.
1995 - Ctenocheles sp. in De Angeli; p. 10, Text-fig. 2 (3 a-c), PI. 1
(fig. 3)
1998 - Ctenocheles sp. in Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 15, Text-figs. 5, 6 (1)
2000 - Ctenocheles sp. in Beschin, De Angeli & Alberti; p. 15
2001 - Ctenocheles sp. in De Angeli & Beschin; p. 10
2006 - Ctenocheles sp. in Beschin, De Angeli, Checchi & Mietto;
p. 99
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: De Angeli (1995) reported one specimen
(MCZ 1490) from “Fontanella” di Grancona (Vicenza);
Beschin et al. (1998) reported one specimen (MCZ
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
17
1484) from “Rossi” quarry of Monte di Malo (Vicenza);
Beschin et al. (2006) reported one specimen (MCZ 2429)
from Buso della Rana (Vicenza).
Family Laomediidae Borrodaile, 1903
Jaxea Nardo, 1847
Type species: Jaxea nocturna Nardo, 1847.
Stratigraphic range: Miocene - Recent.
Jaxea cfr. J. nocturna Nardo, 1 847
1988a - Jaxea cfr. nocturna Nardo in Delle Cave; p. 3, PI. 1 (figs. 1-3),
PI. 2(figs. 1-3)
1993 - Jaxea cfr. nocturna Nardo in Miiller; p. 5, Text-fig. 3 C, D
1998 - Jaxea cfr. nocturna Nardo in Mayoral, Miiller & Muniz; p. 507,
Text-fig. 2 (2)
Stratigraphic range: lower Pliocene.
Occurrence: Toscana.
Material: Delle Cave (1988a) reported ten specimens
(IGF 1018E, 1019E, 1022E, 1033E, 1624E, 1629E,
1632E, 2441 E, 2442E, 2476E) from Castelfiorentino
(Firenze).
Family Upogebiidae Borradaile, 1903
Upogebia Leach, 1814
Type species: Gebia stellata Montagu, 1808.
Stratigraphic range: Jurassic - Recent.
Upogebia perarolensis De Angeli & Messina, 1992
1992 - Upogebia perarolensis De Angeli & Messina; p. 185, Text-
fig. 1, PI. 1 (figs. 1-2); PI. 2 (figs. 1-2)
2001 - Upogebia perarolensis De Angeli & Messina in De Angeli &
Beschin; p. 11, Text-figs. 5- 6
2006 - Upogebia perarolensis De Angeli & Messina in De Angeli &
Rossi; p. 90, PI. 2 (figs. 1, 3)
Holotype: MCZ 1363 (I.G. 284580).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli & Messina (1992) reported 12
specimens (MCZ from 1363 to 1374) from Perarolo
(Vicenza).
Upogebia cfr. U. stellata (Montagu, 1808)
1 89 1 b — Upogebia cfr. stellata (Montagu) in Ristori; p. 24
1914 - Gebia cfr. stellata (Montagu) in M. Gemmellaro; p. 93, PI. 1
(figs. 29-30)
1929 - Upogebia cfr. U. stellata (Montagu) in Glaessner; p. 392
Stratigraphic range: Pliocene - upper Pleistocene
(Sicilian).
Occurrence: Lazio, Sicilia.
Material: Ristori ( 1 89 1 b) reported three specimens
from Farnesina (Roma), housed in Clerici’s collec-
tion, today lost; M. Gemmellaro (1914) reported some
specimens from Ficarazzi (Palermo) (MGUP, catalogue
number unknown) and in thè Marchese di Monterosato’s
collection (repository and catalogue number unknown).
Family Axiidae Huxley, 1879
Etallonia Oppel, 1861
Type species: Maglia longimana Mimster, 1839.
Stratigraphic range: Lower Jurassic (Toarcian) -
Upper Jurassic (Tithonian).
Etallonia sp.
2001 - Etallonia sp. in Garassino & Teruzzi; p. 191, Text-fig. 4
Stratigraphic range: Lower Jurassic (Toarcian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (2001) reported one
specimen (MSNM il 0855) from Sogno (Bergamo).
Note: Garassino & Schweigert (2006) suggested that
this specimen probably could be ascribed to Megachela
Schweigert, 2003.
Huxley caris Bravi & Garassino, 1998
Type species: Huxleycaris beneventana Bravi & Ga¬
rassino, 1998.
Stratigraphic range: Lower Cretaceous (Albian).
Huxleycaris beneventana Bravi & Garassino, 1998
1998a - Huxleycaris beneventana Bravi & Garassino; p. 157, Text-
figs. 26-29
Holotype: M 20885 (N ° 1).
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998a) reported 12
specimens (M 20885: this catalogue number indicates
some specimens, preserved on thè same layer surface;
thè best specimens were marked by thè following Arabie
numbers, 1, 2, 3, 4, 5, 9) from Pietraroia (Benevento).
Protaxius Beurlen, 1930
Type species: Callianassa isochela Woodward, 1876.
Stratygraphic range: Upper Jurassic - middle
Eocene.
Protaxius eocenicus Secretan, 1975
1975 - Protaxius eocenicus Secretan; p. 343, PI. 16 (figs. 1-2, 5)
Holotype: MSNV 4-5 (part and counter-part).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported one specimen
(MSNV 4-5) from Monte Bolca (Verona).
Protaxius sp.
1975 - Protaxius sp. in Secretan; p. 344
18
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported two specimens
(MGPD 10905, 12454) from Monte Bolca (Verona).
Indeterminate family
Orhomalus Etallon, 1861
Type species: Orhomalus virguliuus Etallon, 1861.
Stratigraphic range: Middle Jurassic - ?Lower Cre-
taceous.
Orhomalus rotulensis De Gregorio, 1884
1884 - Orhomalus rotulensis De Gregorio; p. 134
1 929 - Orhomalus rotulensis De Gregorio in Glaessner; p. 283
Holotype: unknown.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: De Gregorio (1884) reported one specimen
from Contrada Rotoli (Palermo) (repository and catalogue
number unknown).
Infraorder Palinura Latreille, 1803
tFamily Coleiidae Van Straelen, 1924
Coleia Broderip, 1835
Type species: Coleia antiqua Broderip, 1835.
Stratigraphic range: Upper Triassic (Camian) -
Lower Cretaceous (Neocomian).
Coleia boboi Garassino & Gironi, 2006
2006 - Coleia boboi Garassino & Gironi; p. 95, Text-figs. 2-6
Holotype: MFSN 20911 a-b.
Stratigraphic range: Upper Triassic (Rhaetian).
Occurrence: Fiurli- Venezia Giulia.
Material: Garassino & Gironi (2006) reported 47
specimens (MFSN: from 28996 to 29014, from 29016 to
29021, from 29023 to 29031, from 29033 to 29045) from
Monte Verzegnis (Udine).
Coleia mediterranea Pinna, 1968
1968 - Coleia mediterranea Pinna; p. 123, Text-figs. 9-10, Pls. 13-16
1990 - Coleia mediterranea Pinna in Teruzzi; p. 96, Text-figs. 11-15
Holotype: MSNM T56.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Pinna (1968) reported one specimen
(MSNM T 56) from Osteno (Como); Teruzzi (1990)
reported five specimens (MSNM Ì7656, Ì7661, Ì7665,
Ì7690, Ì7781) from Osteno (Como).
Coleia pinnai Teruzzi, 1 990
1990 - Coleia pinnai Teruzzi; p. 86, Text-figs. 1-6
Holotype: MSNM i61 1 1 .
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Teruzzi (1990) reported four specimens
(MSNM i6 1 1 1 , Ì7657, Ì7659, Ì7661) from Osteno
(Como).
Coleia popeyei Teruzzi, 1990
1990 - Coleia popeyei Teruzzi; p. 92, Text-figs. 7-10
Holotype: MSNM Ì7683.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Teruzzi (1990) reported seven specimens
(MSNM Ì3369, Ì7683, Ì7684, Ì7686, Ì7687, Ì7688, Ì7780)
from Osteno (Como).
Coleia viallii Pinna, 1 968
1968 - Coleia viallii Pinna; p. 112, Text-figs. 5-8, PI. 3 (fig. 1),
Pls. 4-12
1969 - Coleia viallii Pinna in Pinna; p. 627, Text-figs. 1-4
1990 - Coleia viallii Pinna in Teruzzi; p. 100, Text-figs. 16-18
Holotype: MSNM T51.
Stratigraphic range: Lower Jurassic (Sinemurian).
Occurrence: Lombardia.
Material: Pinna (1968) reported six specimens
(MSNM from T51 to T55, T57) from Osteno (Como);
Pinna (1969) reported two specimens (MSNM i46,
i47) from Osteno (Como); Teruzzi (1990) reported 46
specimens (thè best specimens are: MSNM Ì7604, Ì7658,
Ì7704, Ì7708, Ì7709) from Osteno (Como).
Proeryon Beurlen, 1928
Type species: Eryon hartmanni v. Meyer, 1835.
Stratigraphic range: Lower Jurassic (Toarcian).
Proeryon hartmanni (v. Meyer, 1835)
1835 - Eryon hartmanni v. Meyer; p. 329
1836 - Eryon hartmanni v. Meyer; p. 263, PI. 11 (fig. 1), PI. 12
(figs. 2, 4)
1857 - Eryon hartmanni v. Meyer in Quenstedt; p. 241, PI. 34 (fig. 6)
1 862 - Eryon hartmanni v. Meyer in Oppel; p. 1 1
1 89 1 - Eryon hartmanni v. Meyer in Krause; p. 1 83
1891 - Coleia rnacrophthalma Krause; p. 177, PI. 1 1 (figs. 1-4), nov. syn.
1908 - Eryon hartmanni v. Meyer in Engel; p. 270
1 9 1 1 - Eryon richardsoni Woodward; p. 309, Text-fig. 2, nov. syn.
1925 - Coleia hartmanni (v. Meyer) in Van Straelen; p. 141
1925 - Coleia richardsoni (Woodward) in Woods; p. 22, PI. 6 (fig. 2)
1928 - Proryon hartmanni (v. Meyer) in Beurlen; p. 193, Text-fig. 20,
PI. 6 (fig. 6)
1928 - Proeryon macrophthalmus (Krause) in Beurlen; p. 194, Text-
fig. 21
1928 - Proeryon longiceps Beurlen; p. 196, Text-fig. 22, nov. syn.
1928 - Coleia n. sp. in Beurlen; p. 191
1929 - Proeryon hartmanni (v. Meyer) in Glaessner; p. 339
1929 - Proeryon longiceps Beurlen in Glaessner; p. 339
1929 - Proeryon macrophthalmus (Krause) in Glaessner; p. 340
1944 - Proeryon longiceps Beurlen in Beurlen; p. 374, PI. 34 (fig. 1)
1944 - Proeryon n. sp.? in Beurlen; p. 377, PI. 34 (fig. 2)
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
1952 - Proeryort banzensis Kuhn; p. 156, Text-fig. 1, PI. 13 (figs. 1-2),
PI. 14 (fig. 1), PI. 15 (fig. 1), PI. 16 (fig.l), nov. syn.
1953 - Proeryort hartmanni (v. Meyer) in Hauff; p. 27, PI. 62 a
1969 - Proeryon hartmanni (v. Meyer) in Glaessner; p. 470, Text-fig. 274
1980 - Proeryon hartmanni (v. Meyer) in Forster; p. 76
1980 - Proeryon banzensis Kuhn in Forster; p. 76
1986 - Proeryon macrophthalmus (Krause) in Mudlos; p. 1 50, Text-fig.
without number
1993 -Proeryon sp. in Schmidt-Kaler, Tischlinger & Werner; Text-fig. 47
1993 - Proeryon longiceps Beurlen in Jàger; Text-fig. 47
1998 - Proeryon macrophthalmus (Krause) in Bòttcher; p. 88, Text-
fig. 7.9
2000 - Proeryon hartmanni (v. Meyer) in Schweigert; Text-fig. 2
2001 - Coleia cfr. C. banzensis Kuhn in Garassino & Teruzzi; p. 190,
Text-fig. 3
2001 - Proeryon hartmanni (v. Meyer) in Schweigert; Text-figs. 1-3
200 1 - Proeryon hartmanni (v. Meyer) in Schweigert; Text-fig. 1
2001 - Proeryon sp. in Mauser; p. 106, Text-fig. 4
2005 - Proeryon hartmanni (v. Meyer) in Garassino & Gironi; p. 56,
Text-figs. 4-9
Stratigraphic range: Lower Jurassic (Toarcian).
Occurrence: Lombardia.
Material: Garassino & Teruzzi (2001) reported 15
specimens (MSNM from il 0845 to i 1 0850, i 1 0853,
il 0854, from il 0857 to il 0862, il 0865) from Sogno (Ber¬
gamo); Garassino & Gironi (2005) reported 21 specimens
(MSNM from Ì26209 to i 26212, from Ì26214 to Ì26218
a-b, Ì26220, Ì26222, Ì26223, from Ì26227 to Ì26229 a-b,
from Ì26232 to Ì26236, i 26284 a-b) from Cesana Bri-
anza-Suello (Lecco).
Note: this species is also reported from thè Lower
Jurassic (Toarcian) of Germany.
Pseudocoleia Garassino & Teruzzi, 1993
Type species: Pseudocoleia mazzolenii Garassino &
Teruzzi, 1993.
Stratigraphic range: Upper Triassic (Norian).
Pseudocoleia mazzolenii Garassino & Teruzzi, 1993
1993 - Pseudocoleia mazzolenii Garassino & Teruzzi; p. 19, Text-
figs. 35-40, PI. 4 (figs. 3-4)
1996 - Pseudocoleia mazzolenii Garassino & Teruzzi in Garassino,
Teruzzi & Dalla Vecchia; p. 41
Holotype: MSNM il 2467.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia, Friuli-Venezia Giulia.
Material: Garassino & Teruzzi (1993) reported 36
specimens (thè best specimens are: MSNM il 2467;
MSNB 7560, from 8184 to 8186, 8188) from Ponte
Giurino (Bergamo); Garassino et al. (1996) reported six
specimens (MFSN 1949, 1950, 6153 a-b, 16113 a-b,
16121, 16175) from Caprizzi (Udine).
fFamily Eryonidae De Haan, 1841
Rosenfeldia Garassino, Teruzzi & Dalla Vecchia, 1996
Type species: Rosenfeldia trias ica Garassino, Teruzzi
& Dalla Vecchia, 1996.
Stratigraphic range: Upper Triassic (Norian) - Upper
Jurassic (Tithonian).
Rosenfeldia triasica Garassino, Teruzzi & Dalla Vecchia,
1996
1996 - Rosenfeldia triasica Garassino, Teruzzi & Dalla Vecchia;
p. 33, Text-figs. 6-10, 17-20
Holotype: MFSN 16178.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Friuli-Venezia Giulia.
Material: Garassino et al. (1996) reported 32 speci¬
mens (MFSN 1955, 16110, 16122, 16139, 16141 a-b,
16152, 16155 a-b, from 16171 to 16174 a-b, from 16176
to 16181, 16183, from 16185 to 19197) from Monte Auda
and Forni di Sopra (Udine).
Family Palinuridae Latreille, 1802
Archaeopalinurus Pinna, 1 974
Type species: Archaeopalinurus levis Pinna, 1974.
Stratigraphic range: Upper Triassic (Norian) -
Lower Jurassic (Toarcian).
Archaeopalinurus levis Pinna, 1974
1974 - Archaeopalinurus levis Pinna; p. 29, PI. 14 (fig. 1), PI. 15
(fig. 3), PI. 16 (figs. 4-5)
197 6 - Archaeopalinurus levis Pinna in Pinna; p. 37
1993 - Palinurina cfr. P. longipes Miinster in Dalla Vecchia; p. 65,
Text-fig. 5
1993 - Archaeopalinurus levis Pinna in Garassino & Teruzzi; p. 17,
Text-figs. 29-34, PI. 4 (figs. 1-2)
1996 - Archaeopalinurus levis Pinna in Garassino, Teruzzi & Dalla
Vecchia; p. 42, Text-figs. 21-22
2005 - Archaeopalinurus cfr. A. levis Pinna in Garassino & Gironi;
p. 63, Text-figs. 10-11
Holotype: MSNB 3100.
Stratigraphic range: Upper Triassic (Norian).
Occurrence: Lombardia, Friuli-Venezia Giulia,
Lazio.
Material: Pinna (1974) reported some specimens
(MSNB) from Cene (Bergamo); Pinna (1976) reported
14 specimens (thè best specimens are: MSNBS 2777,
3010) from Rest (Brescia); Garassino & Teruzzi (1993)
reported eight specimens (MSNB 7569 a-b, 7567-7566,
8352; MSNM il 0747, il 0866, il 0740) from Ponte Giurino
(Bergamo); Garassino et al. (1996) reported 126 specimens
(thè best specimens are: MFSN 1392-1412, 1935-1948,
5690, 16108, 16111, 16112, 16115-16120, 16123-16138,
16143, 16144, 16147-16151, 16153, 16154, 16156-16160,
16162-16164) from Monte Auda and Rio Seazza (Udine);
Garassino & Gironi (2005) reported one specimen (MSNM
Ì26237) from Cesana Brianza-Suello (Lecco).
Note: Dalla Vecchia (1993) reported one specimen
(MFSN 15099) from Filettino (Lazio). Even though thè
author ascribed in dubitative form thè specimen to Pali¬
nurina cfr. P. longipes , it could be ascribed to Archaeo¬
palinurus for some morphological characters, typical of
this genus.
20
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Justitia Holthuis, 1946
Type species: Palinurus longimanus H. Milne
Edwards, 1837.
Stratigraphic range: middle Eocene (Lutetian) -
Recent.
Justitia desmaresti (Massalongo, 1854)
1 822 - Palinurus Weber in Desmarest; p. 1 3 1
1840-1850? - Palinurus desmaresti De Zigno; nomen nudum
1 854 - Palinurus Weber in Catullo
1854 - Palinurus desmaresti Massalongo; PI. 12
1855 - Palinurus desmaresti De Zigno in Massalongo; p. 52
1915 - Palinurus desmaresti De Zigno in Fabiani; p. 286
1975 - Palinurus desmaresti De Zigno in Secretan; p. 339, PI. 12
(fig. 1), PI. 13 (figs. 1-5), PI. 14 (figs. 1-4), PI. 15 (figs. 1-4),
PI. 16 (figs. 3-4)
2001 - Justitia desmaresti (Massalongo) in Garassino & Novati; p. 259,
Text-figs. 3-7
2003 - Justitia desmaresti (Massalongo) in Garassino & De Angeli;
p. 131, Text-fig. 1 A
Holotype: MSNV 91-91 bis.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Garassino & Novati (2001) reported 22
specimens (MSNV: 17-17 bis, CR 17, 18, M02, 19, 20,
23B, 23-90 bis, 24, 25-25 bis, 91-91 bis, 92, 93, 94, 95;
MGPD: 6804Z, 7747C-7450C, 7448C-7449C; MSNM:
Ì22867; MFB: I.G. 91130, 1.G. 132590-132605; NHMW:
1 853/XXVII/59- 1 853/XXVII/60) from Monte Bolca
(Verona).
Justitia vicetina Beschin, De Angeli & Garassino, 2001
2001 - Justitia vicetina Beschin, De Angeli & Garassino; p. 91, Text-
figs. 2-4
2001 - Justitia vicetina Beschin, De Angeli & Garassino in Garassino
& Novati; p. 262
2001 - Justitia vicetina Beschin, De Angeli & Garassino in De Angeli
& Beschin; p. 11, Text-fig. 7
2003 - Justitia vicetina Beschin, De Angeli & Garassino in Garassino
& De Angeli; p. 131, Text-fig. 1 B
Holotype: MCZ 1543 (I.G. 284621).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported two
specimens (MCZ 1543, 1544) from “Albanello” quarry
(Vicenza).
Palinurus Weber, 1795
Type species: Astacus elephas Fabricius, 1787, by
monotypy.
Stratigraphic range: Lower Cretaceous (Barremian)-
Recent.
Palinurus sp.
1 998b - Palinurus sp. in Bravi & Garassino; p. 110, Text-fig. 14
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998b) reported
three specimens (M from 21837 to 21839) from Petina
(Salerno).
Palinurus sp.
2000b - Palinurus sp in Garassino; p. 67, Text-fig. 1
Stratigraphic range: Lower Cretaceous (Barremian-
Aptian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino (2000b) reported one
specimen (MFSN 25000) from Torrente Cornappo
(Udine).
Family Scyllaridae Latreille, 1825
Parribacus Dana, 1 852
Type species: Scyllarus antarcticus Lund, 1793.
Stratigraphic range: middle Eocene (Lutetian) -
Recent.
Parribacus cristatus Forster, 1984
1984 - Parribacus cristatus Forster; p. 62, Text-fig. 2
Holotype: MNHB MB. A. 88.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Forster (1984) reported one specimen
(MNHB MB. A. 88) from Monte Bolca (Verona).
Infraorder Anomura MacLeay, 1838
Family Chirostylidae Ortmann, 1892
Eumunida Smith, 1883
Type species: Eumunida pietà Smith, 1883.
Stratigraphic range: upper Eocene (Priabonian) -
Recent.
Eumunida pentacantha (Miiller & Collins, 1991)
1991 - Protomunida pentacantha Miiller & Collins; p. 57, Text-
fig. 2 i, PI. 1 (figs. 15-16)
2000 - Eumunida pentacantha (Miiller & Collins) in Schweitzer &
Feldmann; p. 159
2002 - Eumunida pentacantha (Miiller & Collins) in De Angeli &
Garassino; p. 17, Text-fig. 14, PI. 6 (figs. 3-4)
2003 - Eumunida pentacantha (Miiller & Collins) in De Angeli &
Garassino; p. 99, Text-fig. 1(11)
Holotype: MAFI EBM-1.1 (M. 91-116).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence. Veneto.
Material: De Angeli & Garassino (2002) reported 18
specimens (MSNM from Ì25208 to Ì25218; MCZ from
2167 to 2173) from S. Feliciano (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
21
Family Galatheidae Samouelle, 1819
Calteagalathea De Angeli & Garassino, 2006
Type species: Calteagalathea friulana De Angeli &
Garassino, 2006.
Stratigraphic range: Upper Cretaceous (Maastrich-
tian).
Calteagalathea friulana De Angeli & Garassino, 2006
2001 - Galathea weinfurteri Bachmayer in Beschin, De Angeli &
Checchi; p. 29
2001 - Galathea weinfurteri Bachmayer in De Angeli & Beschin;
p. 11
2002 - Galathea valmaranensis De Angeli & Garassino; p. 8, Text-
fig. 6, PI. 1 (figs. 3-4), PI. 2 (fig. 1)
2003 - Galathea valmaranensis De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1 (4)
2006 - Galathea valmaranensis De Angeli & Garassino in De Angeli &
Rossi; p. 90, PI. 2 (fig. 4)
2006a - Calteagalathea friulana De Angeli & Garassino; p. 274,
| Text-fig. 4 (a-b)
Holotype: MFSN 19969.
Stratigraphic range: Upper Cretaceous (Maastrich-
tian).
Occurrence: Friuli-Venezia Giulia.
Material: De Angeli & Garassino (2006a) reported
two specimens (MFSN 19965, 19969) from Val Caltea
(Pordenone).
Galathea Fabricius, 1793
Type species: Cancer strigosus Linnaeus, 1761.
Stratigraphic range: Cretaceous - Recent.
Galathea affinis Ristori, 1886
1886 - Galathea affinis Ristori; p. 126, PI. 2 (fig. 18)
1909 - Galathea affinis Ristori in Lòrenthey; p. 228
1929 - Galathea affinis Ristori in Glaessner; p. 172
Holotype: unknown.
Stratigraphic range: Miocene, late Pliocene (Piacen-
tian), upper Pleistocene (Sicilian).
Occurrence: Sicilia, Sardegna.
Material: Ristori (1886) reported one specimen
from Bianchi (Palermo) (repository and catalogue
number unknown); Lòrenthey (1909) reported some
specimens from Capo S. Marco (Oristano), housed in
Lovisato’s collection (repository and catalogue number
unknown).
Holotype: MCZ 2228 (I.G. 296470).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
17 specimens (MSNM from i25 147 to i25 1 59; MCZ
2228, 2230, 2231, 2233) from Valmarana (Vicenza); De
Angeli & Messina (1997) and De Angeli & Rossi (2006)
reported two specimens (MCZ 1550, 1551) from Perarolo
(Vicenza).
Galathea cfr. G. weinfurteri Bachmayer, 1950
2002 - Galathea cfr. G. weinfurteri Bachmayer in De Angeli & Garas¬
sino; p. 10, Text-fig. 7, PI. 2 (figs. 2-3)
2003 - Galathea cfr. G. weinfurteri Bachmayer in De Angeli & Garas¬
sino; p. 99, Text-fig. 1 (7)
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002)
reported six specimens (MSNM from Ì25236 to
Ì25238; MCZ 2229, 2232, 2234) from Valmarana
(Vicenza).
Acanthogalathea Miiller & Collins, 1991
Type species: Galathea ( Acanthogalathea ) parva
Miiller & Collins, 1991.
Stratigraphic range: upper Eocene (Priabonian).
Acanthogalathea feldmanni De Angeli & Garassino,
2002
Galathea berica De Angeli & Garassino, 2002
2002 - Galathea berica De Angeli & Garassino; p. 7, Text-fig. 5,
PI. 1 (figs. 1-2)
2003 - Galathea berica De Angeli & Garassino in De Angeli & Garas¬
sino; p. 99, Text-fig. 1(1)
Holotype: MCZ 2186 (I.G. 296428).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
two specimens (MCZ 2186, 2187) from S. Feliciano
(Vicenza).
Galathea valmaranensis De Angeli & Garassino, 2002
1994 - Galathea weinfurteri Bachmayer in Vicariotto & Beschin; p. 7,
PI. 1 (% 1)
1997 - Galathea weinfurteri Bachmayer in De Angeli & Messina;
p. 1 8, Text-fig. 2
2002 - Acanthogalathea feldmanni De Angeli & Garassino; p. 12,
Text-fig. 9, PI. 3 (figs. 3-4)
2003 - Acanthogalathea feldmanni De Angeli & Garassino in
De Angeli & Garassino; p. 99, Text-fig. 1 (9)
Holotype: MCZ 2178.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
two specimens (MCZ 2178, 2188) from S. Feliciano
(Vicenza).
Acanthogalathea parva (Miiller & Collins, 1991)
1991 - Galathea (. Acanthogalathea ) parva Miiller & Collins; p. 56,
Text-fig. 2 h, PI. 2 (fig. 3)
2002 - Acanthogalathea parva (Miiller & Collins) in De Angeli & Ga¬
rassino; p. 11, Text-fig. 8, PI. 2 (fig. 4), PI. 3 (figs. 1-2)
2003 - Acanthogalathea parva (Miiller & Collins) in De Angeli &
Garassino; p. 99, Text-fig. 1 (6)
22
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Holotype: MAFI E.F. 31 (M. 91-106).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002)
reported eight specimens (MSNM from Ì25239 to
Ì25242; MCZ from 2174 to 2177) from S. Feliciano
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Lessinigalathea De Angeli & Garassino, 2002
Type species: Lessinigalathea regale De Angeli &
Garassino, 2002.
Stratigraphic range: lower Eocene (Ypresian).
Lessinigalathea regale De Angeli & Garassino, 2002
2000 - Galathea sp. in Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 8, PI. 1 (fig. 4)
2002 - Lessinigalathaea regale De Angeli & Garassino; p. 13, Text-
fig. 10, PI. 4 (fig. 1)
2003 - Lessinigalathea regale De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1 (2)
Holotype: MCZ 2246 (I.G. 296488).
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Veneto.
Material: Beschin et al. (2000) reported one speci¬
mens (MCZ 1 898) from “Gecchelina” quarry of Monte di
Malo (Vicenza); De Angeli & Garassino (2002) reported
three specimens (MCZ 2246, 2256, 2257) from Monte
Magrè of Schio (Vicenza).
Note: thè correct originai spelling is “ Lessinigalathea ”
instead of “ Lessinigalathaea ” used incorrectly in thè
institution of thè new genus, as misprint.
Palaeomunida Lòrenthey, 1 90 1
Type species: Palaeomunida defecta Lòrenthey,
1901.
Stratigraphic range: Eocene - Oligocene.
Palaeomunida defecta Lòrenthey, 1 90 1
1901 - Palaeomunida defecta Lòrenthey; p. 807, PI. 1 (fig. 3)
1 929 - Palaeomunida defecta Lòrenthey in Lòrenthey & Beurlen;
p. 80, PI. 3 (figs. 3-5)
1933 - Palaeomunida defecta Lòrenthey in Di Salvo; p. 8, PI. 2
(fig. 2 a-d)
1969 - Palaeomunida defecta Lòrenthey in Via Boada; p. 405
1975 - Galathea sp. in Muller; p. 516, 520
1991 - Galathea ( Palaeomunida ) defecta Lòrenthey in Muller & Col¬
lins; p. 56, Text-fig. 2 g, PI. 1 (figs. 12-13), PI. 2 (fig. 1)
2000 - Palaeomunida defecta Lòrenthey in Schweitzer & Feldmann;
p. 158
2001 - Palaeomunida defecta Lòrenthey in Beschin, De Angeli &
Checchi; p. 15, PI. 1 (figs. 2-3)
2002 - Palaeomunida defecta Lòrenthey in De Angeli & Garassino;
p. 14, Text-fig. 1 1, PI. 4 (figs. 2-5), PI. 5 (fig. 1)
2003 Palaeomunida defecta Lòrenthey in De Angeli & Garassino;
p. 99, Text-fig. 1 (3)
Holotype: MAFI.
Stratigraphic range: upper Eocene (Priabonian) -
early Oligocene.
Occurrence: Veneto.
Material: Di Salvo (1933) reported one specimen
from Balzo del Gatto (Palermo) (MGUP, catalogue
number unknown); Beschin et al. (2001) reported 16
specimens (MCZ 2068, 2069, and from 2075 to 2088)
from Castelgomberto (Vicenza); De Angeli & Garassino
(2002) reported nine specimens (MSNM from i25 1 99
to Ì25203, Ì25205, Ì25206; MCZ 2185, 2245) from S.
Feliciano (Vicenza), 15 specimens (MSNM from i25 1 89
to Ì25198, Ì25204, Ì25207; MCZ from 2247 to 2249)
from “Alonte” quarry (Vicenza), 41 specimens (MSNM
from Ì25160 to i25 188 and from Ì25219 to Ì25222;
MCZ from 2235 to 2240, 2241, 2242) from Valmarana
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Palaeomunida multicristata De Angeli & Garassino,
2002
2002 - Palaeomunida multicristata De Angeli & Garassino; p. 15,
Text-fig. 15, PI. 5 (figs. 2-4)
2003 - Palaeomunida multicristata De Angeli & Garassino in
De Angeli & Garassino; p. 99, Text-fig. 1 (5)
Holotype: MCZ 2184 (I.G. 296426).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
nine specimens (MCZ from 2179 to 2184 and from 2221
to 2223) from S. Feliciano (Vicenza).
Spathagalathea De Angeli & Garassino, 2002
Type species: Spathagalathea minuta De Angeli &
Garassino, 2002.
Stratigraphic range: upper Eocene (Priabonian).
Spathagalathea minuta De Angeli & Garassino, 2002
2002 - Spathagalathea minuta De Angeli & Garassino; p. 1 7, Text-
fig. 13, PI. 6 (figs. 1-2)
2003 - Spathagalathea minuta De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1 (8)
Holotype: MCZ 2192 (I.G. 296435).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
nine specimens (MCZ from 2189 to 2197, 2218) from S.
Feliciano and “Alonte” quarry (Vicenza).
Indeterminates
Genus et species indeterminate
1998 - Gen. et sp. indet. in Garassino; p. 69, Text-fig. 4
Stratigraphic range: Lower Cretaceous (Barremian
- Aptian).
Occurrence: Friuli Venezia-Giulia.
Material: Garassino (1998) reported one specimen
(MFSN 21533) from Torrente Comappo (Udine).
CATALOG AND BIBLIOGRAPHY OF THE FOSS1L STOMATOPODA AND DEC APODA FROM ITALY
23
Family Porcellanidae Haworth, 1825
Beripetrolisthes De Angeli & Garassino, 2002
u
Type species: Beripetrolisthes nini Ieri De Angeli &
6 Garassino, 2002.
Stratigraphic range: upper Eocene (Priabonian).
;
Beripetrolisthes mulleri De Angeli & Garassino,
1991 - Porcellanidae sp. B in Miiller & Collins; p. 60, Text-fig. 2 n,
PI. 2 (fig. 2)
2002 - Beripetrolisthes mulleri De Angeli & Garassino; p. 19, Text-
fig. 15, PI. 7 (figs. 1-2)
2003 - Beripetrolisthes mulleri De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1(12)
Holotype: MCZ 2216 (I.G. 296458).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
seven specimens (MCZ from 2205 to 2208, 2216, 2217)
from S. Feliciano (Vicenza).
Eopetrolistlies De Angeli & Garassino, 2002
Type species: Petrolisthesl striatissimus Miiller &
Collins, 1991.
Stratigraphic range: upper Eocene (Priabonian).
Eopetrolìsthes striatissimus (Miiller & Collins, 1991)
1991 - Petrolisthesl striatissimus Miiller & Collins; p. 59, Text-
fig. 2 I, PI. 2 (fig. 8)
2001 - Petrolisthesl striatissimus Miiller & Collins in Beschin,
De Angeli & Checchi; p. 17
2002 - Eopetrolìsthes striatissimus (Miiller & Collins) in De Angeli &
Garassino; p. 20, Text-fig. 16, PI. 7 (figs. 3-4)
2003 - Eopetrolisthes striatissimus (Miiller & Collins) in De Angeli &
Garassino; p. 99, Text-fig. 1(13)
Holotype: MAFI EGA-5.1 (M.91-123).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported 20
specimens (MSNM from Ì25223 to Ì25235; MCZ from
2198 to 2203, 2254) from S. Feliciano (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Lobipetrolisthes De Angeli & Garassino, 2002
Type species: Lobipetrolisthes blowi De Angeli &
Garassino, 2002.
Stratigraphic range: upper Eocene (Priabonian).
Lobipetrolisthes blowi De Angeli & Garassino,
2002
2002 - Lobipetrolisthes blowi De Angeli & Garassino; p. 2 1 , Text-fig.
17, PI. 8 (figs. 1-3)
2003 - Lobipetrolisthes blowi De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1(10)
Holotype: MCZ 2226 (I.G. 296468).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported 15
specimens (MSNM from Ì25243 to Ì25249; MCZ 2219,
2220, from 2224 to 2227, 2252, 2253) from S. Feliciano
(Vicenza).
Longoporcellana Miiller & Collins, 1991
Type species: Longoporcellana denticulata Miiller &
Collins, 1991.
Stratigraphic range: upper Eocene (Priabonian).
Longoporcellana lobata De Angeli & Garassino, 2002
2002 - Longoporcellana lobata De Angeli & Garassino; p. 22, Text-
fig. 18, PI. 8 (fig. 4), PI. 9 (fig. 1)
2003 - Longoporcellana lobata De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1(16)
Holotype: MCZ 2250 (I.G. 296492).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported six
specimens (MCZ from 2212 to 2215, 2250, 2251) from S.
Feliciano (Vicenza).
Petrolisthes Stimpson, 1858
Type species: Porcellana violacea Guérin-Méneville
in Duperry, 1 83 1 , by monotypy.
Stratigraphic range: Eocene - Recent.
Petrolisthes bittneri De Angeli & Garassino, 2002
2002 - Petrolisthes bittneri De Angeli & Garassino; p. 23, Text-
fig. 19, PI. 9 (fig. 2)
2003 - Petrolisthes bittneri De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1 (14)
Holotype: MCZ 2243 (I.G. 296485).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
two specimens (MCZ 2243, 2255) from S. Feliciano
(Vicenza).
Petrolisthes vicetinus Beschin, De Angeli & Checchi,
2001
2001 - Petrolisthes vicetinus Beschin, De Angeli & Checchi; p. 16,
Text-fig. 2, PI. 1 (figs. 1-4)
2002 - Petrolisthes vicetinus Beschin, De Angeli & Checchi in
De Angeli & Garassino; p. 24, Text-fig. 20, PI. 9 (fig. 3)
2003 - Petrolisthes vicetinus Beschin, De Angeli & Checchi in
De Angeli & Garassino; p. 99, Text-fig. 1(15)
Holoty pe: MCZ 2070 (I.G. 286425).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported four
specimens (MCZ 2070, 2093, 2094, 2120) from Cas-
telgomberto (Vicenza); De Angeli & Garassino (2002)
24
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
reported one specimen (MCZ 2244) from Creazzo
(Vicenza).
Pisidia Leach, 1 820
Type species: Cancer longicornis Linnaeus, 1767.
Stratigraphic range: Eocene - Recent.
Pisidia dorsosinuosa De Angeli & Garassino, 2002
2002 - Pisidia dorsosinuosa De Angeli & Garassino; p. 24, Text-
fig. 21, PI. 9 (fig. 4)
2003 - Pisidia dorsosinuosa De Angeli & Garassino in De Angeli &
Garassino; p. 99, Text-fig. 1 (17)
Holotype: MCZ 2210 (I.G. 296452).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2002) reported
three specimens (MCZ from 2209 to 2211) from S. Feli-
ciano (Vicenza).
Superfamily Hippoidea Latreille, 1825
Family Albuneidae Stimpson, 1858
Albunea Weber, 1795
Type species: Albunea symmysta Linnaeus, 1758.
Stratigraphic range: Eocene - Recent.
Albunea cuisiana Beschin & De Angeli, 1984
1984 - Albunea cuisiana Beschin & De Angeli; p. 97, PI. 1 (figs. 1,
1 a), PI. 2 (figs. 1, 1 a-b)
1998 - Albunea cuisiana Beschin & De Angeli in De Angeli; p. 17
2001 - Albunea cuisiana Beschin & De Angeli in De Angeli & Beschin;
p. 12, Text-fig. 9
2005 - Albunea cuisiana Beschin & De Angeli in De Angeli, Beschin
& Checchi; p. 75, PI. 1 (fig. 5)
Holotype: MSNVE 10439.
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Veneto.
Material: Beschin & De Angeli (1984) reported three
specimens (MCZ 1332, 1532; MSNVE 10439) from
“Lovara” and “Boschetto” quarries of Chiampo (Vicenza).
Italialbunea Boyko, 2002
Type species: Albunea lutetiana Beschin & De
Angeli, 1984.
Stratigraphic range: Eocene.
Italialbunea lutetiana (Beschin & De Angeli, 1984)
1984 - Albunea lutetiana Beschin & De Angeli; p. 99, PI. 1 (figs. 2,
2 a), PI. 2 (figs. 2-3, 3 a)
1998 - Albunea lutetiana Beschin & De Angeli in De Angeli; p. 19,
PI. 1 (figs. 1-4)
2001 - Albunea lutetiana Beschin & De Angeli in De Angeli &
Beschin; p. 12
2002 - Italialbunea lutetiana (Beschin & De Angeli) in Boyko; p. 22 1 ,
Text-fig. 72
2004 - Italialbunea lutetiana (Beschin & De Angeli) in Beschin, Busu-
lini. De Angeli & Tessier; p. 1 1 3
2005 - Italialbunea lutetiana (Beschin & De Angeli) in De Angeli,
Beschin & Checchi; p. 75, PI. 1 (figs. 1-3, 3 a)
Holotype: MSNVE 10440.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin & De Angeli (1984) reported one
specimen (MSNVE 10440) from “Main” quarry of Arzi-
gnano (Vicenza); De Angeli (1998) and De Angeli et al.
(2005) reported six specimens (MCZ 1159, 1177, 1375,
1533, 1534, 2403, 1276) from “Main” quarry of Arzig-
nano and “Albanello” quarry of Nogarole Vicentino,
and one specimen (MCZ 1545) from “Alonte” quarry
(Vicenza).
Stemonopa Efford & Haig, 1968
Type species: Stemonopa insignis Efford & Haig,
1968.
Stratigraphic range: Eocene - Recent.
Stemonopa prisca De Angeli, Beschin & Checchi,
2005
2005 - Stemonopa prisca De Angeli, Beschin & Checchi; p.75, Text-
fig. 2, PI. 1 (fig. 4 a-b)
Holotype: MCZ 2404 (I.G. 305 1 1 1 ).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: De Angeli et al. (2005) reported one
specimen (MCZ 2404) from “Main” quarry of Arzignano
(Vicenza).
Zygopa Holthuis, 1961
Type species: Zygopa michaelis Holthuis, 1961.
Stratigraphic range: Oligocene - Recent.
Zygopa galantensis (De Angeli & Marangon, 2001)
2001 - Paralbunea galantensis De Angeli & Marangon; p. 102, Text-
figs. 3-4
2003a - Zygopa galantensis (De Angeli & Marangon) in De Angeli &
Marangon; p. 103, Text-fig. 1 (4 a-b)
2003b - Zygopa galantensis (De Angeli & Marangon) in De Angeli &
Marangon; p. 187, Text-fig. 2 a-b
2004 - Harryhausenia galantensis (De Angeli & Marangon) in Boyko;
p. 935, Text-fig. 1 c
2005 - Zygopa galantensis (De Angeli & Marangon) in De Angeli,
Beschin & Checchi; p. 89
Holotype: MCZ 2061 (I.G. 286339).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte.
Material: De Angeli & Marangon (2001) reported
one specimen (MCZ 2061) from Galanti (Alessan¬
dria).
Note: De Angeli & Marangon (2003 a, b) included
Paralbunea galantensis in Zygopa ; even though
Boyko (2004) suggested to include this species in thè
new genus Harryhausenia, De Angeli & Marangon
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
25
(2003a, b) pointed out that thè morphological char-
acters of thè fossil species are thè same of thè Recent
Zygopa.
Superfamily Paguroidea Latreille, 1802
Family Diogenidae Ortmann, 1 892
Calcinus Dana, 1851
Type species: Calcinus tibicen (Herbst, 1791).
Stratigraphic range: Eocene - Recent.
Calcinus agnoensis Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 - Calcinus agnoensis Beschin, De Angeli, Checchi & Zaran¬
tonello; p. 12, Text-fig. 8, PI. 2 (figs. 5 a-c, 6)
Holotype: MCZ 2356 (I.G. 296575).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported six speci¬
men (MCZ 2284, 2339, 2342, 2355, 2356, 2357) from
Grola di Comedo Vicentino (Vicenza).
Calcinus sp.
1 89 lb - Calcinus sp. in Ristori; p. 24
Stratigraphic range: Pliocene.
Occurrence: Lazio.
Material: Ristori (1891 b) reported two speci¬
men from Monte Mario (Roma), housed in Zuc-
cari’s collection (repository and catalogue number
unknown).
Dardanus Paulson, 1875
Type species: Dardanus hellerii Paulson, 1875.
Stratigraphic range: Eocene - Recent.
Dardanus mediterraneus (Lòrenthey, 1909)
1909 - Pagurus mediterraneus Lòrenthey; p. 226, PI. 2 (fig. 5 a-b)
1929 - Pagurus mediterraneus Lòrenthey in Glaessner; p. 288
2002 - Dardanus mediterraneus (Lòrenthey) in Beschin, Busulini,
De Angeli & Tessier; p. 12
Holotype: unknown.
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported some speci¬
men from Monte della Pace (Cagliari), housed in
Lovisato’s collection (repository and catalogue number
unknown).
Dardanus sp.
2000 - Dardanus sp. in Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 8, PI. 1 (fig. 3)
2001 - Dardanus sp. in De Angeli & Beschin; p. 13
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Veneto.
Material: Beschin et al. (2000) reported one specimen
(MCZ 2013) from “Gecchelina” quarry of Monte di Malo
(Vicenza).
Dardanus sp.
2005 - Dardanus sp. in Beschin, De Angeli, Checchi & Zarantonello;
p. 14, Text-fig. 10, PI. 2 (fig. 3)
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported one speci¬
men (MCZ 2384) from Grola di Comedo Vicentino
(Vicenza).
Diogenes Dana 1851
Type species: Pagurus miles Fabricius, 1787.
Stratigraphic range: Eocene - Recent.
Diogenes sp.
2000 - Diogenes sp. in Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 8, PI. 1 (fig. 2)
2001 - Diogenes sp. in De Angeli & Beschin; p. 13
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Veneto.
Material: Beschin et al. (2000) reported one specimen
(MCZ 1920) from “Gecchelina” quarry of Monte di Malo
(Vicenza).
Eocalcinus \ ia Boada, 1959
Type species: Eocalcinus eocenicus Via Boada,
1959.
Stratigraphic range: Eocene.
Eocalcinus cavits Beschin, Busulini, De Angeli &
Tessier, 2002
1997 - Petrochirus priscus in Vicariotto, p. 27, Text-fig. 1 a-d
2000 - Pagurus sp. in De Angeli & Franchi; p. 20
2001 - Pagurus sp. in De Angeli & Beschin; p. 13
2002 - Eocalcinus cavus Beschin, Busulini, De Angeli & Tessier;
p. 10, Text-fig 5, PI. 1 (fig. 5 a-b), PI. 2 (fig. 1)
2004 - Eocalcinus cavus Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 13
2005 - Eocalcinus cavus Beschin, Busulini, De Angeli & Tessier
in Beschin, De Angeli, Checchi & Zarantonello; p. 12, PI. 2
(fig- 2)
Holotype: MCZ 2263 (I.G. 296391).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Vicariotto (1997) reported one specimen
(MCZ 1574) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza); Beschin et al. (2002) reported two
specimens (MCZ 2263, 2264) from “Main” quarry of
Arzignano (Vicenza); Beschin et al. (2005) reported one
specimen (MCZ 2383) from Grola di Comedo Vicentino
(Vicenza).
26
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Eocalcinus eocenicus Via Boada, 1 959
1959 - Eocalcinus eocenicus V ia Boada; p. 32, Text-fig. 5
1969 - Eocalcinus eocenicus Via Boada in Via Boada; p. 93, Text-
fig. 9, PI. 3 (figs. 1-3)
1989 - Eocalcinus eocenicus Via Boada in Solè & Via Boada;
p. 28
1994 - Eocalcinus eocenicus Via Boada in Beschin, Busulini,
De Angeli & Tessier; p. 164, PI. 1 (fig. 2)
2004 - Eocalcinus eocenicus Via Boada in Beschin, Busulini,
De Angeli & Tessier; p. 1 13
2005 — Eocalcinus eocenicus Via Boada in Beschin, De Angeli, Chec¬
chi & Zarantonello; p. 13
Holotype: MMGB 6649.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported three speci-
mens (MCZ 1436, 1437, 1438) from “Boschetto” quarry
of Nogarole Vicentino (Vicenza).
Note: this species is also reported from thè Eocene of
Spain.
Paguristes Dana, 1851
Type species: Paguristes hirtus Stimpson, 1858.
Stratigraphic range: Cretaceous - Recent.
Paguristes lineatuberculatus Beschin, De Angeli,
Checchi & Mietto, 2006
2006 - Paguristes lineatuberculatus Beschin, De Angeli, Checchi &
Mietto; p. 102, Text-fig. 5, PI. 1 (fig. 8 a-b)
Holotype: MCZ 2432.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (2006) reported one specimen
(MCZ 2432) from Grotta della Poscola (Vicenza).
Paguristes prealpinus Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 - Paguristes prealpinus Beschin, De Angeli, Checchi & Zaran¬
tonello; p. 13, Text-fig. 9, PI. 2 (fig. 4 a-b)
Holotype: MCZ 2340 (LG. 296559).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported four speci-
mens (MCZ 2330, 2331, 2340, 2358) from Grola di
Comedo Vicentino (Vicenza).
Petrochirus Stimpson, 1 859
Type species: Pagurus granulatus Olivieri, 1811.
Stratigraphic range: Eocene - Recent.
Petrochirus mezi (Lòrenthey, 1909)
1909 - Pagurus mezi Lòrenthey; p. 113, PI. 2 (fig. 4 a-b)
1994 - Pagurus cfr. mezi Lòrenthey in Beschin, Busulini, De Angeli &
Tessier; p. 163, PI. 1 (fig. 1 a-c)
2001 - Pagurus cfr. mezi Lòrenthey in De Angeli & Beschin; p. 13
2002 - Petrochirus mezi (Lòrenthey) in Beschin, Busulini, De Angeli &
Tessier; p. 1 1, Text-fig. 6, PI. 1 (fig. 6 a-b)
2004 - Pagurus mezi Lòrenthey in Beschin, Busulini, De Angeli &
Tessier; p. 1 13
2006 - Petrochirus mezi (Lòrenthey) in Beschin, De Angeli, Checchi &
Mietto; p. 99, PI. 1 (fig. 9 a-b)
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported one speci¬
men (MCZ 1313) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza); Beschin et al. (2002) reported one
specimen (MCZ 2265) from “Main” quarry of Arzignano
(Vicenza); Beschin et al. (2006) reported one specimen
(MCZ 2430) from Rio Poscola (Vicenza).
Note: this species is also reported from thè middle
Eocene (Lutetian) of Egypt.
Petrochirus poscolensis Beschin, De Angeli, Checchi &
Mietto, 2006
2006 - Petrochirus poscolensis Beschin, De Angeli, Checchi &
Mietto; p. 100, Text-fig. 4, PI. 1 (fig. 7 a-c)
Holotype: MCZ 2431.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (2006) reported one specimen
(MCZ 243 1 ) from Rio Poscola (Vicenza).
Petrochirus sp.
1995 - Petrochirus sp. in De Angeli; p. 11, Text-fig. 2 (2 a-b), PI. 1
(fig. 4 a-c)
2001 - Petrochirus sp. in De Angeli & Beschin; p. 13
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli (1995) reported two specimens
(MCZ 1491, 1492) from “Fontanella” di Grancona (Vi¬
cenza).
Family Paguridae Latreille, 1802
Anapagurus Henderson, 1888
Type species: Pagurus laevis Bell, 1845.
Stratigraphic range: Pliocene - Recent.
Anapagurus sp.
2004a - Anapagurus sp. in Garassino & De Angeli; p. 37, Text-fig. 3 (9)
Stratigraphic range: Pleistocene.
Occurrence: Emilia Romagna.
Material: Garassino & De Angeli (2004a) reported
one specimen (MG 0635) from Fiume Arda (Piacenza).
Pagurus Fabricius, 1775
Type species: Cancer bernhardus Linnaeus, 1758.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
27
Stratigraphic range: Cretaceous - Recent.
Pagurus arrosor (Herbst, 1794)
1794 - Cancer arrosor Herbst; p. 170, PI. 43 (fig. 1)
1 89 1 b — Pagurus striatus Latreille in Ristori; p. 23
1929 - Pagurus arrosor (Herbst) in Glaessner; p. 287
Holotype: unknown.
Stratigraphic range: Pliocene.
Occurrence: Lazio.
Material: Ristori ( 1 89 1 b) reported some specimens
from Monte Mario (Roma), housed in Rigacci’s and
Clerici’s collections (repository and catalogue number
unknown).
Pagurus manzonii (Ristori, 1888)
1888 -Xanthol manzonii Ristori; p. 213, PI. 4 (figs. 1-4)
1895 -Xanthol manzonii Ristori in Crema; p. 677
1896 - Pagurus manzonii (Ristori) in Ristori; p. 511, PI. 12 (figs. 6-8)
1909 - Pagurus manzonii (Ristori) in Lòrenthey; p. 223
1929 - Pagurus manzonii (Ristori) in Glaessner; p. 287
1950 - Pagurus manzoni (Ristori) in Comaschi Caria; p. 326
1956 - Pagurus manzoni (Ristori) in Comaschi Caria; p. 288
1981 - Pagurus manzonii (Ristori) in Delle Cave; p. 44
Syntypes: IGF 851 E.
Stratigraphic range: Miocene.
Occurrence: Emilia Romagna, Sardegna.
Material: Ristori (1888) reported portions of some
specimens (IGF) from S. Maria Vigiliana (Bologna),
housed in Manzoni ’s collection, and from Monte S.
Michele and Tramezzano (Cagliari), housed in Lovisato’s
collection (repository and catalogue number unknown);
Crema (1895) reported many specimens from Torino
(repository and catalogue number unknown); Ristori
(1896) reported some specimens from Tramezzario di
S. Avendrace and Monte della Pace (Cagliari) (reposi¬
tory and catalogue number unknown); Lòrenthey (1909)
reported one specimen from S. Avendrace (Cagliari),
housed in Lovisato’s collection (repository and catalogue
number unknown).
Pagurus squamosus Ristori, 1886
1886 - Pagurus squamosus Ristori; p. 125, PI. 3 (figs. 3-5)
1929 -Pagurus? ( Calcinus ?) squamosus Ristori in Glaessner; p. 288
1981 - Pagurus squamosus Ristori in Delle Cave; p. 44
Holotype: IGF 860E.
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (1886) reported one specimen (IGF
860E) from Sarteano (Siena).
Pagurus substriatus A. Milne Edwards, 1861
1846 - Pagurus striatus Latreille in E. Sismonda; p. 70, PI. 3 (fig. 8)
1861 - Pagurus substriatus A. Milne Edwards in E. Sismonda;
p. 20
1886 - Pagurus substriatus A. Milne Edwards in Ristori; p. 124, PI. 3
(figs. 14-15)
1929 - Pagurus substriatus A. Milne Edwards in Glaessner; p. 288
Holotype: lost.
Stratigraphic range: Pliocene.
Occurrence: Piemonte, Toscana, Sardegna.
Material: A. Milne Edwards (1861) reported one
specimen from Asti, today lost; E. Sismonda (1846)
reported one specimen from Asti, today lost; E. Sismonda
(1861) reported this species without pointing out number
of specimens and repository; Ristori (1886) reported two
specimens from Volterra (Firenze), today lost.
Note: E. Sismonda (1846) reported one specimen
ascribing it to Pagurus striatus , later E. Sismonda (1861)
ascribed thè same specimen to Pagurus substriatus.
Pagurus cfr. P substriatus A. Milne Edwards, 1861
1896 - Pagurus cfr. substriatus A. Milne Edwards in Ristori; p. 510,
PI. 12 (fig. 5)
1909 - Pagurus cfr. substriatus A. Milne Edwards in Lòrenthey;
p. 225
1929 - Pagurus cfr. substriatus A. Milne Edwards in Glaessner; p. 288
1950 - Pagurus cfr. substriatus A. Milne Edwards in Comaschi Caria;
p. 326
1956 - Pagurus cfr. substriatus A. Milne Edwards in Comaschi Caria;
p. 288
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Ristori (1896) reported one specimen from
Capo S. Elia (Cagliari) (repository and catalogue number
unknown); Lòrenthey (1909) reported two specimens
from Capo S. Elia and Monte S. Michele (Cagliari)
(repository and catalogue number unknown).
Pagurus sp.
1889 -Pagurus sp. in Ristori; p. 408, PI. 15 (figs. 14-16)
1929 - Pagurus sp. in Glaessner; p. 289
1974 -Pagurus sp. in Mastrorilli; p. 4
Stratigraphic range: Miocene.
Occurrence: Liguria.
Material: Ristori (1889) reported some specimens
from Dego and Sassello (Savona), housed in don Perran-
do’s collection, today lost and in Michelotti’s collection
(repository and catalogue number unknown).
Pagurus sp.
1902 - Pagurus sp. in Lovisato; p. 8
1950 - Pagurus sp. in Comaschi Caria; p. 326
1956 -Pagurus sp. in Comaschi Caria; p. 288
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lovisato (1902) reported some specimens
from S. Bartolomeo (Cagliari) (repository and catalogue
number unknown).
Pagurus sp.
1914 - Pagurus sp. in M. Gemmellaro; p. 92
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
28
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: M. Gemmellaro (1914) reported some
specimens from Ficarazzi (Palermo) (MGUP, catalogue
number unknown).
Pagurus sp.
2004a - Pagurus sp. in Garassino & De Angeli; p. 37, Text-fig. 3 (4-6)
Stratigraphic range: Pleistocene.
Occurrence: Emilia Romagna.
Material: Garassino & De Angeli (2004a) reported
three specimens (MG from 0632 to 0634) from Fiume
Arda (Piacenza).
Pagurus sp.
2004 - Pagurus sp. in Garassino, De Angeli, Gallo & Pasini; p. 256,
Text-fig. 2 a-c
Stratigraphic range: Pliocene.
Occurrence: Piemonte.
Material: Garassino et al. (2004) reported four speci¬
mens (PU from 41144 to 41147) from Masserano and
Cossato (Biella).
Indeterminate
Genus indeterminate
1995 - Genus indet. in Collins & Dieni; p. 69, Text-fig. 2 (3, 5, 6).
Stratigraphic range: Upper Cretaceous (Cenoma-
nian).
Occurrence: Friuli-Venezia Giulia.
Material: Collins & Dieni (1995) reported one speci¬
men (MGPD 242) from Col dei Schiosi (Pordenone).
Infraorder Brachyura Latreille, 1 802
Section Podotremata Guinot, 1977
Subsection Dromiacea De Haan, 1833
Superfamily Homolodromioidea Alcock, 1899
tFamily Prosopidae v. Meyer, 1860
Pithonoton v. Meyer, 1 842
Type species: Prosopon marginatum v. Meyer, 1842.
Stratigraphic range: Jurassic - Paleocene.
Pithonoton bidentatum (Reuss, 1858)
1858 - Goniodromites bidentatus Reuss; p. 12
1869 - Prosopon etalloni G. G. Gemmellaro; p. 12, PI. 2 (figs. 50-51),
nov. syn.
1929 - Pithonoton ( Goniodromites ) etalloni (G. G. Gemmellaro) in
Glaessner; p. 327
1988 - Pithonoton bidentatum (Reuss) in Wehner; p. 87, PI. 6
(Figs. 4-6)
1998 - Pithonoton bidentatum (Reuss) in Miiller; p. 1 7
Holotype: lost.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: G. G. Gemmellaro (1869) reported some
specimens from Castello di Termini (Palermo), housed
in Battagliai collection (MGUP, catalogue number
unknown).
Note: this species is also reported from thè Jurassic of
Germany and Austria.
Pithonoton grande (v. Meyer, 1 860)
1860 - Prosopon grande v. Meyer; p. 202, PI. 23 (figs. 10-13)
1869 - Prosopon polyphemi G. G. Gemmellaro; p. 17, PI. 2 (fig. 59),
nov. syn.
1925 - Pithonoton polyphemi (G. G. Gemmellaro) in Van Straelen;
p. 367
1929 - Pithonoton polyphemi (G. G. Gemmellaro) in Glaessner;
' p. 323
1988 - Pithonoton grande (v. Meyer) in Wehner; p. 91, PI. 7 (figs. 1-2)
1998 - Pithonoton grande (v. Meyer) in Miiller; p. 17
Lectotype: BSPG 1881 IX 678.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: G. G. Gemmellaro (1869) reported one
specimen from Villabate (Palermo) (MGUP, catalogue
number unknown).
Note: this species is also reported from thè Jurassic of
Austria.
Pithonoton marginatum (v. Meyer, 1 842)
1842 - Prosopon marginatum v. Meyer; p. 72, PI. 15 (fig. 3)
1869 - Prosopon marginatum v. Meyer in G. G. Gemmellaro; p. 1 1,
PI. 2 (figs. 48-49)
1929 - Pithonoton marginatum (v. Meyer) in Glaessner; p. 322
1988 - Pithonoton marginatum (v. Meyer) in Wehner; p. 79, PI. 5
(fig. 8), PI. 6 (fig. 1)
1998 - Pithonoton marginatum (v. Meyer) in Miiller; p. 18
2000 - Pithonoton marginatum (v. Meyer) in Miiller, Krobicki &
Wehner; p. 54, Text-figs. 12, 17 K, L, 18 A-E
2006a - Pithonoton marginatum (v. Meyer) in De Angeli & Garassino;
p. 277 , Text-fig. 5
Neotype: BSPG 1881 1X 681.
Stratigraphic range: Upper Jurassic (Oxfordian -
Tithonian).
Occurrence: Friuli-Venezia Giulia, Sicilia.
Material: G. G. Gemmellaro (1869) reported two
specimens from Castello di Termini and Villabate (Pa¬
lermo), housed in Battaglia’s collection (MGUP, cata¬
logue number unknown); De Angeli & Garassino (2006a)
reported one specimen (MFSN 15763) from Altopiano
Prat (Udine).
Note: this species is also reported from thè Jurassic of
Germany and Austria.
Indeterminate family
Oxythyreus Reuss, 1858
Type species: Oxythyreus gibbus Reuss, 1858.
Stratigraphic range: Upper Jurassic (Tithonian).
Oxythyreus gibbus Reuss, 1858
1858 - Oxythyreus gibbus Reuss; p. 12
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
29
1 859 - Oxythyreus gibbus Reuss in Reuss; p. 75, PI. 24 (figs. 8-9)
1 860 - Oxythyreus gibbus Reuss in v. Meyer; p. 2 1 8
1869 - Oxythyreus gibbus Reuss in G. G. Gemmellaro; p. 18
1889 - Oxythyreus gibbus Reuss in Moericke; p. 56
1895 - Oxythyreus gibbus Reuss in Remes; p. 201, PI. 1 (fig. 5)
1905 - Oxythyreus gibbus Reuss in Remes; p. 35
1929 - Oxythyreus gibbus Reuss in Glaessner; p. 285
1969 - Oxythyreus gibbus Reuss in Glaessner; p. 489, Text-fig. 301 (2)
Holotype: unknown.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: G. G. Gemmellaro (1869) reported one
specimen from Misilmeri (Palermo) (MGUP, catalogue
number unknown).
Note: this species is also reported from thè Upper
Jurassic (Tithonian) of Moravia.
Superfamily Dromioidea De Haan, 1833
Family Dromiidae De Haan, 1833
Dromia Weber, 1795
Type species: Cancer personatus Linnaeus, 1758.
Stratigraphic range: Eocene - Recent.
Dromia vulgaris H. Milne Edwards, 1837
1837 - Dromia vulgaris H. Milne Edwards; p. 173
1 885 - Dromia vulgaris H. Milne Edwards in Carus; p. 498
1914 - Dromia vulgaris H. Milne Edwards in M. Gemmellaro; p. 77,
PI. 1 (fig- 1)
1929 - Dromia vulgaris H. Milne Edwards in Glaessner; p. 139
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported one speci¬
men fforn Monte Pellegrino (Palermo) (MGUP, catalogue
number unknown).
IDromia sp.
1933 - ? Dromia sp. in Di Salvo; p. 1 1, PI. 2 (fig. 7 a-b)
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported one specimen
from La Pietra Lunga (Palermo) (MGUP, catalogue
number unknown).
Noetlingia Beurlen, 1928
Type species: Dromia claudiopolitana Bittner, 1893.
Stratigraphic range: Eocene - Oligocene.
Noetlingia claudiopolitana (Bittner, 1 893)
1893 - Dromia claudiopolitana Bittner; p. 21, PI. 2 (fig. 5)
1928 - Noetlingia claudiopolitana (Bittner) in Beurlen; p. 166
1929 - Noetlingia claudiopolitana (Bittner) in Glaessner; p. 274
1929 - Noetlingia claudiopolitana (Bittner) in Lòrenthey & Beurlen;
p. 99, PI. 4 (figs. 8-9)
1994 - Noetlingia claudiopolitana (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 166, PI. 1 (fig. 4)
2001 - Noetlingia claudiopolitana (Bittner) in De Angeli & Beschin;
p. 14
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported two speci-
mens (MCZ 1212, 1252) from “Boschetto” quarry of
Nogarole Vicentino and “Lovara” quarry of Chiampo
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Noetlingia veronensis (Bittner, 1886)
1886 -Dromia veronensis Bittner; p. 46, PI. 1 (fig. 2)
1901 - Dromia veronensis Bittner in Oppenheim; p. 284
1915 - Dromia veronensis Bittner in Fabiani; p. 285
1928 - Noetlingia veronensis (Bittner) in Beurlen; p. 166
1929 - Noetlingia veronensis (Bittner) in Glaessner; p. 274
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Bittner (1883) reported one specimen from
San Giovanni in Valle (Verona) (repository and catalogue
number unknown).
Family Dynomenidae Ortmann, 1892
Basinotopus McCoy, 1 849
Type species: Inachus lamarckii Desmarest, 1822.
Stratigraphic range: Eocene.
Basinotopus lamarcki (Desmarest, 1 822)
1822 - Inachus Lamarcki Desmarest; p. 116, PI. 9 (figs. 15-16)
1849 - Basinotopus Lamarckii (Desmarest) in McCoy; p. 168
1854 - Basinotopus Lamarckii (Desmarest) in McCoy; p. 123
1858 - Dromilites Lamarckii (Desmarest) in Bell; p. 29, PI. 5
(figs. 1-9)
1898 - Dromilites Lamarcki (Desmarest) in Carter; p. 19
1928 - Dromilites lamarcki (Desmarest) in Beurlen; p. 163
1933 - Dromia Fabr.l (sic!) (Desmarest) in Di Salvo; p. 11, PI. 2
(fig- 7)
1983 - Dromilites lamarcki (Desmarest) in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 57, PI. 1 (fig. 2)
2001 - Dromilites lamarcki (Desmarest) in De Angeli & Beschin;
p. 14
2002 - Basinotopus lamarckii (Desmarest) in Collins; p. 83
2004 - Basinotopus lamarcki (Desmarest) in Beschin, Busulini,
De Angeli & Tessier; p. 1 13
2005 - Basinotopus lamarcki (Desmarest) in Beschin, De Angeli,
Checchi & Zarantonello; p. 15, PI. 2 (fig. 9)
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto, Sicilia.
30
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: Di Salvo (1933) reported one specimen
from La Pietra Lunga (Palermo) (catalogue number
unknown); Busulini et al. (1983) reported one speci¬
men (*11) from “Main” quarry of Arzignano (Vicenza);
Beschin et al. (2005) reported one specimen (MCZ 2338)
from Grola di Comedo Vicentino (Vicenza).
Note: this species is also reported from thè lower
Eocene (Ypresian) of Great Britain.
Cyamocarcinus Bittner, 1883
Type species: Cyamocarcinus angustifrons Bittner,
1883.
Stratigraphic range: lower/upper Eocene.
Cyamocarcinus angustifrons Bittner, 1883
1883 - Cyamocarcinus angustifrons Bittner; p. 310, PI. 1 (fig. 8)
1894 - Cyamocarcinus angustifrons Bittner in De Gregorio; p. 9,
PI. 1 (fig- 2)
1897 - Cyamocarcinus angustifrons Bittner in Lòrenthey; p. 155
1898 - Cyamocarcinus angustifrons Bittner in Lòrenthey; p. 53, PI. 3
(fig- 2)
1899 - Cyamocarcinus budensis Oppenheim; p. 58; nov. syn.
1905 - Cyamocarcinus angustifrons Bittner in Checchia-Rispoli;
p. 314, PI. 1 (figs. 5-8)
191 Oa — Cyamocarcinus angustifrons Bittner in Fabiani; p. 26
1929 - Cyamocarcinus angustifrons Bittner in Glaessner; p. 131
1929 - Cyamocarcinus angustifrons Bittner in Lòrenthey & Beurlen;
p. 219, PI. 10 (figs. 2, 6)
1933 - Cyamocarcinus angustifrons Bittner in Di Salvo; p. 27
1969 - Cyamocarcinus angustifrons Bittner in Via Boada; p. 372
1975 - Cyamocarcinus angustifrons Bittner in Miiller; p. 516, 520
1991 - Cyamocarcinus angustifrons Bittner in Miiller & Collins; p. 64,
PI. 3 (figs. 9-10)
2000 - Cyamocarcinus angustifrons Bittner in Beschin, Busulini,
De Angeli, Tessier & Ungaro; p. 8, PI. 2 (fig. 1 a-b)
2001 - Cyamocarcinus angustifrons Bittner in De Angeli & Beschin;
p. 14
Holotype: unkonwn.
Stratigraphic range: lower/upper Eocene (Ypresian -
Priabonian).
Occurrence: Veneto, Sicilia.
Material: Bittner (1883) reported one specimen
from Monte Magrè di Schio (Vicenza) (repository
and catalogue number unknown); De Gregorio (1894)
reported one specimen from Monte Postale di Bolca
(Verona) (repository and catalogue number unknown);
Checchia-Rispoli (1905) reported one specimen from
Balzo del Gatto (Palermo) (MGUP, catalogue number
unknown); Di Salvo (1933) reported 30 specimens
from Balzo del Gatto, La Pietra Lunga, and Bichinello
(Palermo) (MGUP, catalogue number unknown);
Beschin et al. (2000) reported one specimen (MCZ
1678) from “Gecchelina” quarry of Monte di Malo
(Vicenza).
Note: this species is also reported from thè upper
Eocene of Hungary.
Cyclothyreus Remes, 1 895
Type species: Cyclothyreus strambergensis Remes,
1895.
Stratigraphic range: Upper Jurassic (Tithonian).
Cyclothyreus oxythyreiforme (G. G. Gemmellaro, 1869)
1869 - Prosopon oxythyreiforme G. G. Gemmellaro; p. 15, PI. 2
(fig. 58), PI. 3 (fig. 1)
1889 - Prosopon oxythyreiforme G. G. Gemmellaro in Moericke;
p. 57, PI. 6 (fig. 10)
1895 - Prosopon oxythyreiforme G. G. Gemmellaro in Remes; p. 202,
PI. 2 (fig. 9)
1 897 - Prosopon cfr. P. oxythyreiforme G. G. Gemmellaro in Roman;
p. 277, PI. 8 (fig. 12)
1911 - Prosopon oxythyreiforme G. G. Gemmellaro in Blaschke;
p. 151
1913 - Prosopon oxythyreiforme G. G. Gemmellaro in Joukowski &
Favre; p. 489, PI. 34 (fig. 8)
1925 - Prosopon oxythyreiforme G. G. Gemmellaro in Van Straelen;
p. 374, Text-fig. 169
1928 - Oxythyreus oxythyreiforme (G. G. Gemmellaro) in Beurlen;
p. 150
1929 - Cyclothyreus oxythyreiforme (G. G. Gemmellaro) in Glaessner;
p. 132
1998 - Cyclothyreus oxythyreiformis (G. G. Gemmellaro) in Miiller;
p. 20
Holotype: unknown.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: G. G. Gemmellaro (1869) reported three
specimens from Villabate and Misilmeri (Palermo)
(MGUP, catalogue number unknown).
Note: this species is also reported from thè Upper
Jurassic (Tithonian) of Germany and Austria, and thè
Lower Cretaceous (Neocomian) of France.
Cyclothyreus reussi (G. G. Gemmellaro, 1869)
1869 - Prosopon reussi G. G. Gemmellaro; p. 13, PI. 2 (figs. 52-54)
1925 - Cyclothyreus reussi (G. G. Gemmellaro) in Van Straelen; p. 376
1929 - Pithonoton reussi (G. G. Gemmellaro) in Glaessner; p. 326
1998 - Cyclothyreus reussi (G. G. Gemmellaro) in Wehner; p. 69
2000 - Cyclothyreus reussi (G. G. Gemmellaro) in Miiller, Krobicki &
Wehner; Text-fig. 2 1
Holotype: unknown.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
Material: G. G. Gemmellaro (1869) reported many
specimens from Villabate (Palermo) (MGUP, catalogue
number unknown).
Cyclothyreus tithonium (G. G. Gemmellaro, 1869)
1869 - Prosopon tìthonium G. G. Gemmellaro; p. 14, PI. 2
(figs. 55-57)
1925 - Cyclothyreus tithonium (G. G. Gemmallaro) in Van Straelen;
p. 377
1929 - Pithonoton tithonium (G. G. Gemmellaro) in Glaessner;
p. 326
1998 - Cyclothyreus tithonium (G. G. Gemmellaro) in Wehner; p. 69
Holotype: unknown.
Stratigraphic range: Upper Jurassic (Tithonian).
Occurrence: Sicilia.
CATALOG AND BIBLIOGRAPHY OF THE FOSS1L STOMATOPODA AND DECAPODA FROM ITALY
31
Material: G. G. Gemmellaro (1869) reported two
specimens from Villabate (Palermo) (MGUP, catalogue
number unknown).
Dromilites H. Milne Edwards, 1837
Type species: Dromia bucklandii H. Milne Edwards,
1837.
Stratigraphic range: Paleocene - Miocene.
Dromilites corvini (Bittner, 1 893)
1893 - Dromia corvinii Bittner; p. 16, PI. 2 (fig. 6)
1 898 - Dromia corvini Bittner in Lòrenthey; p. 1 1 8
1928 - Dromia corvini Bittner in Beurlen; p. 168
1 929 - Dromia corvini Bittner in Glaessner, p. 1 38
1929 - Dromilites corvini (Bittner) in Lòrenthey & Beurlen; p. 98,
PI. 4 (figs. 6-7)
2001 - Dromilites corvini (Bittner) in Beschin, De Angeli & Checchi;
p. 17, Text-fig. 3, PI. 1 (figs. 6-7)
2001 - Dromilites corvini (Bittner) in De Angeli & Beschin; p. 13
Holotype: unknown.
Stratigraphic range: upper Eocene (Priabonian) -
lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported three speci¬
mens (MCZ 2123, 2125, 2126) from Castelgomberto
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Dromilites hilarionis (Bittner, 1883)
1883 - Dromia Hilarionis Bittner; p. 306, PI. 1 (fig. 5)
191 Oa - Dromia Hilarionis Bittner in Fabiani; p. 25
1915 - Dromia Hilarionis Bittner in Fabiani; p. 284
1928 - Pseudodromilites hilarionis (Bittner) in Beurlen; p. 168
1929 - Dromilites hilarionis (Bittner) in Glaessner; p. 140
1983 - Dromilites hilarionis (Bittner) in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 57, PI. 1 (fig. 4)
1989 - Dromilites hilarionis (Bittner) in Solè & Via Boada; p. 28
1995 - Dromilites hilarionis (Bittner) in De Angeli; p. 12, Text-fig. 2
(5), PI. 1 (fig. 6)
2001 - Dromilites hilarionis (Bittner) in De Angeli & Beschin;
p. 13
2004 - Dromilites hilarionis (Bittner) in Beschin, Busulini, De Angeli
& Tessier; p. 1 1 3
2005 - Dromilites hilarionis (Bittner) in Beschin, De Angeli, Checchi
& Zarantonello; p. 14, PI. 2 (fig. 7)
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priaboanian).
Occurrence: Veneto.
Material: Bittner (1883) reported one specimen
from Ciupio (Verona) (repository and catalogue number
unknown); Busulini et al. (1893) reported four specimens
(*5, *7, *134; MSNVE 10090) from “Main” quarry of
Arzignano (Vicenza); De Angeli (1995) reported one
specimen (MCZ 1494) from “Fontanella” di Grancona
(Vicenza); Beschin et al. (2005) reported two specimens
(MCZ 2345; MV 51) from Grola di Comedo Vicentino
(Vicenza).
Note: this species is also reported from thè middle/
upper Eocene (Lutetian - Priabonian) of Spain.
Dromilites pastoris Via Boada, 1959
1959 - Dromilites pastoris Via Boada; p. 364, Text-fig. 6
1969 - Dromilites pastoris Via Boada in Via Boada; p. 99, PI. 4
(fig- 5)
1983 - Dromilites pastoris Via Boada in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 58, PI. 1 (fig. 1 a-b)
1989 - Dromilites pastoris Via Boada in Solè & Via Boada; p. 28
1994 - Dromilites pastoris Via Boada in Beschin, Busulini, De Angeli
& Tessier; p. 165, PI. 1 (fig. 3)
2001 - Dromilites pastoris Via Boada in De Angeli & Beschin;
p. 14
2004 - Dromilites pastoris Via Boada in Beschin, Busulini, De Angeli
& Tessier; p. 113
2005 - Dromilites pastoris Via Boada in Beschin, De Angeli, Checchi
& Zarantonello; p. 15, PI. 2 (fig. 8)
Holotype: MGSB 15.995.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1983) reported two speci¬
mens (*136; MSNVE 10091) from “Main” quarry of
Arzignano (Vicenza); Beschin et al. (1994) reported one
specimen (MCZ 1456) from “Boschetto” quarry ofNoga-
role Vicentino (Vicenza); Beschin et al. (2005) reported
three specimens (MCZ 2285, 2286, 2287) from Grola di
Comedo Vicentino (Vicenza).
Note: this species is also reported from thè Eocene of
Spain.
Dynomene Desmarest, 1 823
Type species: Dynomene ispida Guérin-Méneville,
1832.
Stratigraphic range: Oligocene - Recent.
Dynomene lessinea Beschin, De Angeli & Checchi,
2001
2001 - Dynomene lessinea Beschin, De Angeli & Checchi; p. 17,
Text-fig. 4, PI. 1 (figs. 5, 8)
2001 - Dynomene lessinea Beschin, De Angeli & Checchi in De Angeli
& Beschin; p. 14
Holotype: MCZ 2065 (I.G. 286420).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported five speci¬
mens (MCZ 2065, from 2089 to 2092) from Castelgom¬
berto (Vicenza).
Gemmellarocarcinus Checchia-Rispoli, 1905
Type species: Gemmellarocarcinus lòrenthey i Chec¬
chia-Rispoli, 1905.
Stratigraphic range: middle Eocene (Lutetian).
Gemmellarocarcinus lorentheyi Checchia-Rispoli, 1905
1905 - Gemmellarocarcinus lorentheyi Checchia-Rispoli; p. 316,
PI. 1 (figs. 1-2)
32
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1929 - Gemmellarocarcìnus lòrentheyi Checchia-Rispoli in Glaessner;
p. 182
1933 - Gemmellarocarcìnus lòrentheyi Checchia-Rispoli in Di Salvo;
p. 33
1969 - Gemmellarocarcìnus lòrentheyi Checchia-Rispoli in Via Boada;
p. 379
1975 - Gemmellarocarcìnus lòrentheyi Checchia-Rispoli in Miiller;
p. 510, 516
1991 - Gemmellarocarcìnus lòrentheyi Checchia-Rispoli in Miiller &
Collins; p. 64, PI. 2 (fig. 15)
Holotype: MGUP.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1905) reported one spec¬
imen from Balzo del Gatto (Palermo) (MGUP, catalogue
number unknown); Di Salvo (1933) reported one speci¬
men from Balzo del Gatto (Palermo) (MGUP, catalogue
number unknown).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Etungary.
Graptocarcinus Roemer, 1887
Type species: Graptocarcinus texanus Roemer, 1887.
Stratigraphic range: Lower/Upper Cretaceous.
Graptocarcinus bellonii Collins & Dieni, 1995
1995 - Graptocarcinus bellonii Collins & Dieni; p. 70, Text-figs. 2
(1,2,4)
2006a - Graptocarcinus bellonii Collins & Dieni in De Angeli &
Garassino; p. 279, Text-fig. 6
Holotype: MGPD B241.
Stratigraphic range: Upper Cretaceous (Maastrich-
tian).
Occurrence: Friuli-Venezia Giulia.
Material: Collins & Dieni (1995) reported one speci¬
men (MGPD B241) from Col dei Schiosi (Pordenone);
De Angeli & Garassino (2006a) reported one specimen
(MFSN 16937) from Borgo Vigant (Udine).
Kromtitis Miiller, 1984
Type species: Dromilites koberi Bachmayer & Toll-
mann, 1953.
Stratigraphic range: Eocene - Miocene.
Kromtitis tetratubercu/atus Beschin, Busulini, De Angeli
& Tessier, 2002
2002 - Kromtitis tetratuberculatus Beschin, Busulini, De Angeli &
Tessier; p. 12, Text-fig. 7, PI. 2 (figs. 2-3 a-b)
2004 - Kromtitis tetratuberculatus Beschin, Busulini, De Angeli &
Tessier in Beschin, Busulini, De Angeli & Tessier; p. 1 13
Holotype: MCZ 2266 (EG. 296394).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported three speci-
mens (MCZ 2266, 2267, 2268) from “Main” quarry of
Arzignano (Vicenza).
Palaeodromites A. Milne Edwards, 1865
Type species: Cyphonotus incertus Bell, 1863.
Stratigraphic range: Upper Jurassic - Upper Creta¬
ceous.
Palaeodromites himeraensis (Checchia-Rispoli, 1914)
1914 - Distefania himeraensis Checchia-Rispoli; p. 177, Text-
figs. 1-2., PI. 1 (figs. 2-3)
1914 - Distefania siculo Chechia-Rispoli; p. 181, Text-figs. 3-4, PI. 1
(fig. 1), nov. syn.
1950 - Cyphonotus himeraensis (Checchia-Rispoli) in Wright &
Wrigth; p. 22, Text-fig. 1 1
1972 - Palaeodromites himeraensis (Checchia-Rispoli) in Wright &
Collins; p. 51, Text-fig. 9 (f)
Holotype: unknown.
Stratigraphic range: Upper Cretaceous (Cenoma-
nian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1914) reported two
specimens from Termini Imerese (Palermo) (repository
and catalogue number unknown).
Note: Wright & Collins (1972) considered P. siculo as
synonym with P. himeraensis.
Subsection Archaeobrachyura Guinot, 1977
Superfamily Raninoidea De Haan, 1839
Family Raninidae De Haan, 1839
Cosmonotus Adams & Withe, 1 848
Type species: Cosmonotus grayi Adams & Withe, 1848.
Stratigraphic range: Eocene - Recent.
Cosmonotus eocaenicus Beschin, Busulini, De Angeli &
Tessier, 1988
1988 - Cosmonotus eocaenicus Beschin, Busulini, De Angeli &
Tessier; p. 160, Text-fig. 2, PI. 1 (figs. 1-4)
2001 - Cosmonotus eocaenicus Beschin, Busulini, De Angeli & Tessier
in De Angeli & Beschin; p. 20
2004 - Cosmonotus eocaenicus Beschin, Busulini, De Angeli & Tessier
in Beschin, Busulini, De Angeli & Tessier; p. 113
Holotype: MCZ 1 1 05 (I.G. 2 1 1 644).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) reported four speci¬
mens (MCZ from 1105 to 1108) from “Main” quarry of
Arzignano (Vicenza).
Cyrtorhina Monod, 1 956
Type species: Cyrtorhina granulosa Monod, 1956.
Stratigraphic range: Eocene - Recent.
Cyrtorhina globosa Beschin, Busulini, De Angeli &
Tessier, 1988
1988 - Cyrtorhina globosa Beschin, Busulini, De Angeli & Tessier;
p. 163, Text-fig. 3, PI. 2 (fig. 1 a-d)
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
33
1998 - Cyrtorhina globosa Beschin, Busulini. De Angeli & Tessier in
Rizzotto; p. 21, Pls. 1-2
i 2001 - Cyrtorhina globosa Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 14, Text-figs. 10 (4 a-c), 1 1
; 2004 - Cyrtorhina globosa Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 113
2004 - Cyrtorhina globosa Beschin, Busulini, De Angeli & Tessier
in Tessier, Busulini, Beschin & De Angeli; p. 9, Text-figs. 4 a-c,
5
2005 - Cyrtorhina globosa Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli, Checchi & Zarantonello; p. 16
Holotype: MCZ 1135 (I.G. 211671).
Stratigraphic range: lower/middleEocene(Ypresian-
Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) reported one specimen
(MCZ 1135) from “Boschetto” quarry of Chiampo (Vi¬
cenza); Rizzotto (1998) reported one specimen (MCZ
1614) from “Main” quarry of Arzignano (Vicenza);
Tessier et al. (2004) reported one specimen (MGPD
29012) from Zovo of Bolca (Verona).
1994 - Lianira beschini Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, Busulini, De Angeli & Tessier; p. 175, PI. 4
(figs. 1-2 a-b)
1998 - Lianira beschini Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, Busulini, De Angeli, Tessier & Ungaro; p. 22,
Text-figs. 9(1), 10(1-3)
2000 - Lianira beschini Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, De Angeli & Alberti; p. 15
2001 - Lianira beschini Beschin, Busulini, De Angeli, Tessier &
Ungaro in De Angeli & Beschin; p. 20, Text-fig. 14 (3 a-c)
Holotype: MCZ 1231 (I.G. 211746).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1991, 1994) reported four
specimens (SV 278; MCZ 1231, 1232, 1233) from “Bos¬
chetto” quarry of Nogarole Vicentino (Vicenza); Beschin
et al. (1998) reported one specimen (MCZ 1537) from
“Rossi” quarry of Monte di Malo (Vicenza).
Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro, 1991
Cyrtorhina oblonga Beschin, Busulini, De Angeli &
Tessier, 1988
1988 - Cyrtorhina oblonga Beschin, Busulini, De Angeli & Tessier;
p. 166, Text-fig. 4, PI. 3 (figs. 1-3)
1997 - Cyrtorhina oblonga Beschin, Busulini, De Angeli & Tessier in
Vicariotto; p. 29, Text-fig. 2 a-b
2001 - Cyrtorhina oblonga Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 16, Text-fig. 10 (1 a-b)
2004 - Cyrtorhina oblonga Beschin, Busulini, De Angeli & Tessier in
Tessier, Busulini, Beschin & De Angeli; Text-fig. 7
2004 - Cyrtorhina oblonga Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 13
2005 - Cyrtorhina oblonga Beschin, Busulini, De Angeli & Tessier
in Beschin, De Angeli, Checchi & Zarantonello; p. 15, PI. 3
(fig- 1)
Holotype: MCZ 1100 (I.G. 211639).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) reported six speci¬
mens (MCZ 1100, 1101, 1102, 1103, 1104; SV 230) from
“Main” quarry of Arzignano (Vicenza); Vicariotto (1997)
reported one specimen (MCZ 1577) from “Boschetto”
quarry of Nogarole Vicentino (Vicenza); Beschin et al.
(2005) reported one specimen (MCZ 2288) from Grola di
Comedo Vicentino (Vicenza).
Lianira Beschin, Busulini, De Angeli, Tessier & Ungaro,
1991
Type species: Lianira beschini Beschin, Busulini, De
Angeli, Tessier & Ungaro, 1991.
Stratigraphic range: middle Eocene (Lutetian).
Lianira beschini Beschin, Busulini, De Angeli, Tessier &
Ungaro, 1991
1991 - Lianira beschini Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 197, Text-fig. 4, PI. 1 (fig. 1 a-e), PI. 2 (figs. 1-2 a-c),
PI. 3 (fig. 1 a-d)
1991 - Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 199, Text-fig. 5, PI. 4 (figs. 1 a-b, 3 a-c)
1994 - Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, Busulini, De Angeli & Tessier; p. 176, PI. 4
(fig- 4)
1998 - Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, Busulini, De Angeli, Tessier & Ungaro; p. 22,
figs. 9(4), 10(4-6)
2000 - Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, De Angeli & Alberti; p. 15
2001 - Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro in De Angeli & Beschin; p. 20, Text-fig. 14 (4 a-c)
2004 - Lianira convexa Beschin, Busulini, De Angeli, Tessier &
Ungaro in Beschin, Busulini, De Angeli & Tessier; p. 1 13
Holotype: MCZ 1219 (I.G. 211734).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1991, 1994) reported nine
specimens (MCZ from 1217 to 1222, 1303, 1316, 1317)
from “Main” quarry of Arzignano (Vicenza) and “Bos¬
chetto” quarry of Nogarole Vicentino (Vicenza); Beschin
et al. (1998) reported one specimen (MCZ 1538) from
“Rossi” quarry of Monte di Malo (Vicenza).
Lophoranina Fabiani, 1910
Type species: Ranina marestiana Kònig, 1825.
Stratigraphic range: Upper Cretaceous - Oligocene.
Lophoranina aldrovandii Ronzani, 1818
1 594 - Sepites in Aldrovandi; carte 224, 225, 25 1
1648 - Sepites in Ambrosino; p. 452
1818 - Ranina Aldrovandi Ronzani; p. 76, PI. 5 (figs. 3, 4)
1822 - Ranina Aldrovandi Ronzani in Desmarest: p. 121, PI. 6 (fig. 1),
not PI. 10 (figs. 5-7)
1837 - Ranina Aldrovandi Ronzani in A. Milne Edwards; p. 195
1838 - Ranina Aldrovandi Ronzani in A. Milne Edwards in Lamarck;
p. 195
1846 - Ranina Aldrovandi Ronzani in E. Sismonda; p. 64
34
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1 859 - Ranina Aldrovandi Ronzarli in Reuss; p. 1 9, 20
1861 - Ranina Aldrovandi Ronzani in E. Sismonda; p. 17, non Text-
figs. 16-17
1861 - Hippa Aldrovandi (Ronzani) in A. Milne Edwards in Cuvier;
p. 106
1861 - Ranina aldrovandi Ronzani in Michelotti; p. 141
1866 - Ranina Aldrovandi Ronzani in Woodward; p. 591
1 872 - Ranina Aldrovandi Ronzani in A. Milne Edwards; p. 7
1875 - Ranina Aldrovandi Ronzani in Bittner; p. 4
1 889 - Ranina Aldrovandi Ronzani in Ristori; p. 408
1907 - Ranina Aldrovandi Ronzani in Gortani; p. 10
191 Ob — Ranina ( Lophoranina ) Aldrovandii Ronzani in Fabiani; p. 95,
Text-figs. 1-4
1929 - Ranina {Lophoranina) Aldrovandii Ronzani in Glaessner,
p. 364
1966 - Lophoranina aldrovandi (Ronzani) in Via Boada; p. 254
1969 - Lophoranina aldrovandi (Ronzani) in Via Boada; p. 1 19, 391
1974 - Ranina aldrovandii ? (Ronzani) in Mastrorilli; p. 4
1988 - Lophoranina aldrovandii (Ronzani) in Beschin, Busulini,
De Angeli & Tessier; p. 188
Holotype: MPP Geo 00000046 (ex number
G 1 04 1 5 Rocc/493 3 6)
Stratigraphic range: ?Eocene.
Occurrence: ?NE Italy.
Material: Ronzani (1818), Reuss (1859) and Fabiani
(191 Ob) reported thè species probably from thè Eocene
of N Italy.
Lophoranina bittnerì (Lòrenthey, 1902)
1875 - Ranina nov. spec.? in Bittner; p. 66, PI. 1 (fig. 3)
1 898 - Ranina cfr. Marestiana in Lòrenthey; p. 22
1902 - Ranina Bittnerì Lòrenthey; p. 809, PI. 1 (figs. 1-2)
1905 - Ranina Bittnerì Lòrenthey in Airaghi; p. 203, PI. 4 (fig. 1)
191 Oa — Ranina Bittnerì Lòrenthey in Fabiani, p. 20
191 Ob - Ranina ( Lophoranina ) Bittnerì Lòrenthey in Fabiani; p. 90
1915 - Ranina Bittneri Lòrenthey in Fabiani; p. 284, 285
1 929 - Ranina ( Lophoranina ) Bittneri Lòrenthey in Lòrenthey & Beur-
len; p. 114, PI. 5 (figs. 2-3)
1929 - Ranina ( Lophoranina ) Bittneri Lòrenthey in Glaessner,
p. 365
1933 - Ranina Bittneri Lòrenthey in Di Salvo; p. 13, PI. 1 (fig. 4 a-b)
1966 - Ranina bittneri Lòrenthey in Ancona; p. 406
1983 - Lophoranina bittneri (Lòrenthey) in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 60, PI. 1 (fig. 5)
1988 - Lophoranina bittneri (Lòrenthey) in Beschin, Busulini,
De Angeli & Tessier; p. 179, Text-fig. 7 (3), PI. 6 (figs. 2-4)
1998 - Lophoranina bittneri (Lòrenthey) in Beschin, Busulini,
De Angeli, Tessier & Ungaro; p. 20, Text-fig. 8 (3)
2000 - Lophoranina bittneri (Lòrenthey) in Beschin, De Angeli &
Alberti; p. 15
2001 - Lophoranina bittneri (Lòrenthey) in De Angeli & Beschin,
p. 17, Text-fig. 13 (5)
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto, Sicilia.
Material: Bittner (1875) reported this species from
Monte Zuello, Ciupio (Verona) (repository and catalogue
number unknown); Airaghi (1905) reported one speci¬
men from Lonigo (Vicenza) (MNAV, catalogue number
unknown); Di Salvo (1933) reported one specimen Ponte
di Castronuovo (Palermo) (MGUP, catalogue number
unknown); Busulini et al. (1983) reported one speci¬
men (*141) from “Main” quarry of Arzignano (Vicenza);
Beschin et al. (1988, 1998) reported four specimens
(MSNVE 11877,11 878, MCZ 1117,1118) from Castello
di Soave (Verona) and “Rossi” quarry of Monte di Malo
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Lophoranina laevifrons (Bittner, 1875)
1875 - Ranina laevifrons Bittner; p. 68, PI. 1 (fig. 4)
1895 - Ranina laevifrons Bittner in Bittner; p. 247, PI. 1 (figs. 3-4)
191 Oa — Ranina laevifrons Bittner in Fabiani; p. 20
191 Ob — Ranina {Lophoranina) laevifrons Bittner in Fabiani; p. 90
1915 - Ranina laevifrons Bittner in Fabiani; p. 284
1966 - Ranina laevifrons Bittner in Ancona; p. 406
1983 - Lophoranina laevifrons (Bittner) in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 61, PI. 2 (fig. 4)
1988 - Lophoranina laevifrons (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 181, Text-fig. 7 (4-6), PI. 7 (figs. 3-5)
1994 - Lophoranina laevifrons (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 174, PI. 3 (fig. 3)
2001 - Lophoranina laevifrons (Bittner) in De Angeli & Beschin; p. 18,
Text-fig. 13 (4 a-b)
2004 - Lophoranina laevifrons (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 1 13
2005 - Lophoranina laevifrons (Bittner) in Beschin, De Angeli, Chec¬
chi & Zarantonello; p. 16, PI. 3 (fig. 3)
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875, 1895) reported two speci¬
mens from S. Giovanni Ilarione (Verona) (repository
and catalogue number unknown); Busulini et al. (1983)
reported one specimen (*36) from “Main” quarry of
Arzignano (Vicenza); Beschin et al. (1988) reported six
specimens (SV 36, 279, 280, 281; MCZ 1115, 1116)
from “Albanello” and “Boschetto” quarries of Nogarole
Vicentino, “Main” quarry of Arzignano (Vicenza), and
Crocegrande (Verona); Beschin et al. (1994) reported
fìve specimens (MCZ 1261, 1262, 1263, 1275, 1435)
from “Albanello” and “Boschetto” quarries of Nogarole
Vicentino (Vicenza); Beschin et al. (2005) reported one
specimen (MCZ 2344) from Grola di Comedo Vicentino
(Vicenza).
Lophoranina marestiana (Kònig, 1 825)
1 8 1 7 - Remipes sulcatus in Desmarest; p. 5 1 2
1 822 - Ranina Aldrovandi in Desmarest; p. 5 12
1825 - Ranina Maresiana Kònig; p. 2, PI. 1 (fig. 15, R. Maretiana)
1854 - Ranina Aldrovandii Ronzani in Catullo; p. 886
1859 - Ranina Marestiana Kònig in Reuss; p. 20, PI. 5 (figs. 1-2)
1872 - Ranina Maresiana {sic!) Kònig in A. Milne Edwards; p. 8
1875 - Ranina Marestiana Kònig in Bittner; p. 64, PI. 1 (figs. 1-2)
1883 - Ranina Marestiana Kònig in Bittner; p. 300, PI. 1 (figs. 1-2)
1908 - Ranina Marestiana Kònig in Fabiani; p. 209, 236
191 Oa — Ranina marestiana Kònig in Fabiani; p. 19
191 Ob — Ranina {Lophoranina) marestiana Kònig in Fabiani; p. 89
1915 - Ranina marestiana Kònig in Fabiani; p. 284, 285
1915 - Ranina marestiana Kònig in Dainelli; p. 699
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
35
1933 - Ranina cfr. Marestiana Kònig in Di Salvo; p. 12
1959 - Lophoranina marestiana (Kònig) in Via Boada; p. 365
1966 - Ranina marestiana Kònig in Ancona; p. 402, Text-figs. 1-2,
Pls. 1-3
1 966 - Lophoranina marestiana (Kònig) in Via Boada; p. 240, Text-
figs. 1-3, PI. 1
1968 - Ranina ( Lophoranina ) marestiana (Kònig) in Vogeltanz; p. 33
1969 - Lophoranina marestiana (Kònig) in Via Boada; p. 104, Text-
fig. 1 1, PI. 5 (fig. 1), PI. 6 (fig. 1)
1988 - Lophoranina marestiana (Kònig) in Beschin, Busulini,
De Angeli & Tessier; p. 175, Text-fig. 6, PI. 5 (figs. 2-4),
PI. 6 (fig. 1 a-c)
1994 - Lophoranina marestiana (Kònig) in Beschin, Busulini,
De Angeli & Tessier; p. 173, PI. 3 (fig. 4)
1998 - Lophoranina marestiana (Kònig) in Beschin Busulini,
De Angeli, Tessier & Ungaro; p. 20, Text-figs. 6 (2-3), 8(1)
1998 - Lophoranina marestiana (Kònig) in Miiller; p. 25
2000 - Lophoranina marestiana (Kònig) in Beschin, De Angeli &
Alberti; p. 15
2001 - Lophoranina marestiana (Kònig) in De Angeli & Beschin;
p. 17, Text-figs. 12-13 (1 a-c)
2006a - Lophoranina marestiana (Kònig) in De Angeli & Garassino;
p. 280, Text-fig. 7 (a-b)
Holotype: unknown.
Stratigraphic range: lower/upper Eocene (Ypresian-
Priabonian).
Occurrence: Veneto, Friuli-Venezia Giulia, Sicilia.
Material: Reuss (1859), A. Milne Edwards (1872),
Bittner (1875, 1883) reported this species from Vicen¬
za and Verona (repository and catalogue number
unknown); Dainelli (1915) reported four specimens
from Buttrio and Valle del Natisone (Pordenone)
(repository and catalogue number unknown); Di Salvo
(1933) reported two specimens from Ponte di Castro¬
nuovo and Torre Malfitano (Palermo) (MGUP, cata¬
logue number unknown); Ancona (1966) reported this
species from “Zanconato” quarry of Chiampo (Vicen¬
za) (MCSM, catalogue number unknown); Beschin et
al. (1988, 1994, 1998) reported 18 specimens (SV 293,
294, 297, 298, 299, 300, 302, 315, 316; MSN VE from
11873 to 11876; MCZ from 1121 to 1125, 1331, 1536)
from Chiampo and “Rossi” quarry of Monte di Malo
(Vicenza); De Angeli & Garassino (2006a) reported
four specimens (MFSN from 29047 to 29050) from
Almadis (Pordenone).
Note: this species is also reported from Eocene of
Spain, Austria, and N Africa.
Lophoranina marestiana var. avesana (Bittner, 1883)
1883 - Ranina Marestiana var. avesana Bittner; p. 301, PI. 1 (fig. 2)
1884 - Ranina Marestiana var. avesana Bittner in Bittner; p. 16, PI. 1
(figs. 1-3)
1929 - Ranina ( Lophoranina ) Marestiana var. avesana Bittner in
Glaessner, p. 367
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1883, 1884) reported seven
specimens from Avesa Valley (Verona), belonging to
Nicolisi’s collection (repository and catalogue number
unknown).
Lophoranina maxima Beschin, Busulini, De Angeli &
Tessier, 2004
1988 - Lophoranina cf. reussi Woodward in Beschin, Busulini,
De Angeli & Tessier; p. 185, Text-fig. 8, PI. 5 (fig. 1), PI. 8
(figs. 1-4), PI. 9 (fig. 1)
2001 - Lophoranina cf. reussi Woodward in De Angeli & Beschin;
p. 18
2004 - Lophoranina maxima Beschin, Busulini, De Angeli & Tes¬
sier; p. 110, Text-figs. 1-2, PI. 1 (figs.1-3), PI. 2 (figs. 1-2)
Holotype: MCZ 1127 (LG. 211663).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988, 2004) reported eight
specimens (SV 37, from 318 to 322; MCZ 1127; MCSM,
catalogue number unknown) from “Main” quarry of
Arzignano (Vicenza).
Lophoranina reussi (Woodward, 1 866)
1859 - Ranina sp. in Reuss; p. 21, PI. 5 (figs. 3-4)
1866 - Ranina Reussii Woodward; p. 591
1883 - Ranina Reussii Woodward in Bittner; p. 302
1898 - Ranina Reussi Woodward in Lòrenthey; p. 18, PI. 2 (fig. 1)
1 899 - Ranina Reussi Woodward in Oppenheim; p. 58
1905 - Ranina Reussi Woodward in Airaghi; p. 203, PI. 4 (fig. 2)
1908 - Ranina Reussi Woodward in Fabiani; p. 210, 236
1 9 1 Oa - Ranina Reussi Woodward in Fabiani; p. 19
191 Ob — Ranina ( Lophoranina ) Reussi Woodward in Fabiani; p. 89
1915 - Ranina Reussi Woodward in Fabiani; p. 284, 285
1 929 - Ranina {Lophoranina) Reussi Woodward in Lòrenthey & Beur-
len; p. Ili, PI. 5 (fig. 1)
1959 - Lophoranina reussi (Woodward) in Via Boada; p. 365
1966 - Lophoranina reussi (Woodward) in Via Boada; p. 246, Text-
fig. 4, PI. 2 (figs. 1-3)
1966 - Ranina reussi Woodward in Ancona; p. 406
1969 - Lophoranina reussi (Woodward) in Via Boada; p. 110, Text-
fig. 12, PI. 5 (fig. 2), PI. 6 (figs. 2-4)
1988 - Lophoranina reussi (Woodward) in Beschin, Busulini,
De Angeli & Tessier; p. 183, Text-figs. 7 (1-2), PI. 7 (figs. 1-2)
1995 - Lophoranina reussi (Woodward) in De Angeli; p. 1 1
2001 - Lophoranina reussi (Woodward) in De Angeli & Beschin; p. 18,
Text-fig. 13 (2 a-b)
2005 - Lophoranina reussi (Woodward) in Beschin, De Angeli, Chec¬
chi & Zarantonello; p. 16, PI. 3 (fig. 2)
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Reuss (1859), Bittner (1883), Airaghi ( 1 905)
reported this species from Lessini orientali and Monti
Berici (repository and catalogue number unknown); Fabia¬
ni (1910a) reported this species from Nanto, Montruglio,
Castello di Barbarano, Villaga, Sarego, Lonigo and
Grancona (Vicenza), and S. Giovanni Barione (Verona)
(repository and catalogue number unknown); Beschin et
al. (1988) reported two specimens (SV 309; MCZ 1119)
from Villaga and Barbarano (Vicenza); De Angeli (1995)
reported one specimen (MCZ 1493) from “Fontanella”
di Grancona (Vicenza); Beschin et al. (2005) reported
three specimens (MCZ 2325, 2326, 2327) from Grola di
Comedo Vicentino (Vicenza).
36
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Note: this species is also reported from thè middle
Eocene (Lutetian) of Spain and upper Eocene (Priaboni-
an) of Hungary.
Lophoranina straeleni Via Boada, 1959
1959 - Lophoranina straeleni Via Boada; p. 366, Text-fig. 7
1969 - Lophoranina straeleni Via Boada in Via Boada; p. 115, Text-
fig. 13, PI. 7 (fig. 1), PI. 8 (fig. 1)
1 989 - Lophoranina straeleni Via Boada in Solè & Via Boada; p. 29
1998 - Lophoranina straeleni Via Boada in Beschin, Busulini,
De Angeli & Tessier; p. 22, Text-figs. 6 (5), 8 (2)
2000 - Lophoranina straeleni Via Boada in Beschin, De Angeli &
Alberti; p. 15
2001 - Lophoranina straeleni Via Boada in De Angeli & Beschin;
p. 18
Holotype: MNCNM 1.001.
Stratigraphic range: lower/middle Eocene (Ypresian-
Lutetian).
Occurrence: Veneto.
Material: Via Boada (1959, 1969) reported three
specimens from Bolca area (Verona) (MS, catalogue
number unknown); Beschin et al. (1998) reported one
specimen (MCZ 1539) from “Rossi” quarry of Monte di
Malo (Vicenza).
Note: this species is also reported from thè middle
Eocene (Lutetian) of Spain.
Lophoranina sp.
1 982 - Lophoranina sp. in Busulini, Tessier & Visentin; p. 78
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1982) reported one speci¬
men from “Main” quarry of Arzignano (Vicenza) (SV,
catalogue number unknown).
Lovarina Beschin, Busulini, De Angeli, Tessier &
Ungaro, 1991
Type species: Lovarina cristata Beschin, Busulini, De
Angeli, Tessier & Ungaro, 1991.
Stratigraphic range: middle Eocene (Lutetian).
Lovarina cristata Beschin, Busulini, De Angeli, Tessier
& Ungaro, 1991
1991 - Lovarina cristata Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 202, Text-fig. 6, PI. 5 (figs. 1 a-b, 3)
2001 - Lovarina cristata Beschin, Busulini, De Angeli, Tessier &
Ungaro in De Angeli & Beschin; p. 20, Text-fig. 14 (5 a-c)
Holotype: MCZ 1224 (I.G. 211739).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1991) reported nine
specimens (MCZ from 1224 to 1230, 1299, 1318) from
“Lovara” quarry of Chiampo (Vicenza).
Ly sirude Goeke, 1985
Type species: Lyreidus tridentatus De Haan, 1841.
Stratigraphic range: Eocene - Recent.
Lysirude paronae (Crema, 1895)
1895 - Lyreidus paronae Crema; p. 671, Text-fig. 11
1 907 - Lyreidus paronae Crema in Sacco; p. 1 16
1929 - Lyreidus paronae Crema in Glaessner; p. 241
1998 - Lysirude paronae (Crema) in Tucker; p. 324
2004 - Lysirude paronae (Crema) in Garassino, De Angeli, Gallo &
Pasini; p. 258, Text-figs. 3-4
Holotype: PU 80111.
Stratigraphic range: middle Miocene (Helvetian) -
Pliocene.
Occurrence: Piemonte.
Material: Crema (1895) reported one specimen (PU
80111), from Sciolze (Torino); Garassino et al. (2004)
reported one specimen (PU 41148) from Orta S. Giulio
(Novara).
Notopoides Henderson, 1888
Type species: Notopoides latus Henderson, 1888.
Stratigraphic range: Eocene - Recent.
Notopoides exiguus Beschin, Busulini, De Angeli &
Tessier, 1988
1988 - Notopoides exiguus Beschin, Busulini, De Angeli & Tessier;
p. 188, Text-fig. 9, PI. 9 (figs. 2-3)
2001 - Notopoides exiguus Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 17, Text-fig. 10 (2)
Holotype: MSNVE 11879.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) reported three speci¬
mens (SV 325 ; MSNVE 11879; MCZ 1128) from “Bos¬
chetto” quarry of Chiampo (Vicenza).
Notopus De Haan, 1 84 1
Type species: Cancer dorsipes Linnaeus, 1758.
Stratigraphic range: Eocene - Recent.
Notopus bey rie hi Bittner, 1875
1875 - Notopus Beyrichii Bittner; p. 72, PI. 1 (fig. 6)
1884 - Notopus Beyrichii Bittner in Bittner; p. 17, PI. 1 (fig. 4)
1 898 - Notopus Beyrichii Bittner in Lòrenthey; p. 26
1 899 - Notopus Beyrichi Bittner in Oppenheim; p. 58
1 9 1 Oa - Notopus Beyrichi Bittner in Fabiani; p. 20, 29, 31
1915 - Notopus Beyrichi Bittner in Fabiani; p. 284
1929 - Notoporanina Beyrichi (Bittner) in Lòrenthey & Beurlen;
p. 117, PI. 5 (fig. 4-6)
1929 - Notoporanina Beyrichi (Bittner) in Glaessner; p. 278
1950 - Notopus beyrichi Bittner in Malaroda; p. 191
1982 - Notopus beyrichi Bittner in Busulini, Tessier & Visentin;
p. 78
1988 - Notopus beyrichi Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 191, Text-fig. 10, PI. 10 (fig. 1 a-b)
2001 - Notopus beyrichi Bittner in De Angeli & Beschin; p. 20, Text-
fig. 14(2 a-b)
2004 - Notopus beyrichi Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 1 13
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
37
2005 - Notopus beyrichi Bittner in Beschin, De Angeli, Checchi &
Zarantonello; p. 16, PI. 3 (fig. 5)
2006 - Notopus beyrichi Bittner in Beschin, De Angeli, Checchi &
Mietto; p. 103, PI. 2 (fig. 1)
Holotype: MNHB K.107, MB.A. 657.
Stratigraphic range: middle/upper Eocene
(Lutetian-Priabonian) and lower Oligocene (Rupe-
lian).
Occurrence: Veneto.
Material: Bittner (1875, 1884) reported this spe-
cies (MNHB K.107, MB.A. 657) from S. Giovanni
Barione and Castelrotto (Verona); Malaroda (1950)
reported one specimen from Montecchio of Costozza
(Vicenza) (repository and catalogue number unknown);
Busulini et al. (1982) reported two specimens from
“Main” quarry of Arzignano (Vicenza) (repository and
catalogue number unknown); Beschin et al. (1988)
reported five specimens (SV 274, 304, 306; MCZ1 1 12,
1113) from “Main” quarry of Arzignano (Vicenza);
Beschin et al. (2005) reported two specimens (MCZ
2359; MCV 50) from Grola di Comedo Vicentino
(Vicenza); Beschin et al. (2006) reported one specimen
(MCZ 2439) from Buso della Rana (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Quasilaeviranina Tucker, 1998
Type species: Ranina simplicissima Bittner, 1883.
Stratigraphic range: Eocene.
Quasilaeviranina arzignanensis (Beschin, Busulini, De
Angeli & Tessier, 1988)
1988 - Notosceles arzignanensis Beschin, Busulini, De Angeli &
Tessier; p. 194, Text-fig. 11, PI. 10 (figs. 2-3)
1998 - Quasilaeviranina arzignanensis (Beschin, Busulini, De Angeli
& Tessier) in Tucker; p. 356
2001 - Quasilaeviranina arzignanensis (Beschin, Busulini, De
Angeli & Tessier) in De Angeli & Beschin; p. 16, Text-fig. 10
(3 a-c)
2004 - Quasilaeviranina arzignanensis (Beschin, Busulini, De Angeli
& Tessier) in Beschin, Busulini, De Angeli & Tessier; p. 1 13
Holotype: MCZ 1114.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) reported two speci¬
mens (SV 276; MCZ 1114) from “Main” quarry of Arzi¬
gnano (Vicenza).
Quasilaeviranina ombonii (Fabiani, 1910)
1910a - Ranina Ombonii Fabiani; p. 2, PI. 2 (fig. 1)
191 Ob - Ranina Ombonii Fabiani in Fabiani; p. 91
1915 - Ranina Ombonii Fabiani in Fabiani; p. 284
1929 - Ranina ( Laeviranina ) Ombonii Fabiani in Glaessner;
p. 364
1988 - Laeviranina ombonii (Fabiani) in Beschin, Busulini, De Angeli
& Tessier; p. 169, Text-fig. 5 (3), PI. 3 (figs. 4-6)
1998 - Quasilaeviranina ombonii (Fabiani) in Tucker; p. 357
2001 - Quasilaeviranina ombonii (Fabiani) in De Angeli & Beschin;
p. 17, Text-fig. 10 (7)
Holotype: MGPD 7820.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Fabiani (191 Ob) reported one speci¬
men (MGPD 7820) from Villaga (Vicenza); Beschin
et al. (1988) reported two specimens (MCZ 1131;
MSNVE 11872) from Nanto (Vicenza) and Breonio
(Verona).
Quasilaeviranina simplicissima (Bittner, 1883)
1883 - Ranina simplicissima Bittner; p. 305, PI. 1 (fig. 4 a-b)
191 Oa - Ranina simplicissima Bittner in Fabiani; p. 3
191 Ob - Ranina simplicissima Bittner in Fabiani; p. 91
1915 - Ranina simplicissima Bittner in Fabiani; p. 284
1929 - Ranina ( Laeviranina ) simplicissima Bittner in Glaessner;
p. 364
1929 - Laeviranina simplicissima (Bittner) in Lorenthey & Beurlen;
p. 106, PI. 4 (fig. 11)
1983 - Laeviranina cfr. simplicissima (Bittner) in Busulini, Tessier,
Visentin, Beschin, De Angeli & Rossi; p. 59, PI. 1 (fig. 3).
1988 - Laeviranina cf. simplicissima (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 173, Text-fig. 5 (1), PI. 4 (figs. 4-5)
1994 - Laeviranina cf. simplicissima (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 173, PI. 3 (fig. 2)
1998 - Quasilaeviranina simplicissima (Bittner) in Tucker; p. 355
2001 - Quasilaeviranina simplicissima (Bittner) in De Angeli &
Beschin; p. 16, Text-fig. 10 (5)
2004 - Quasilaeviranina simplicissima (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 1 1 3
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1883) reported one specimen
from Monte Vegroni of Bolca (Verona) (repository and
catalogue number unknown); Busulini et al. (1983)
reported two specimens (*136; MSNVE 10092) from
“Main” quarry of Arzignano (Vicenza); Beschin et al.
(1988) reported six specimens (SV 136, 326; MSNVE
10092; MCZ from 1132 to 1134,) from “Main” quarry
of Arzignano, “Boschetto” and “Albanello” quarries of
Nogarole Vicentino (Vicenza); Beschin et al. (1994)
reported four specimens (MCZ 1133, 1209, 1210,
1272) from “Boschetto” quarry of Nogarole Vicentino
(Vicenza).
Note: this species is also reported from thè middle
Eocene (Lutetian) of Hungary.
Ranilia H. Milne Edwards, 1837
Type species: Ranilia muricata H. Milne Edwards,
1837.
Stratigraphic range: Eocene - Recent.
Ranilia punctulata Beschin, Busulini, De Angeli &
Tessier, 1988
1988 - Ranilia punctulata Beschin, Busulini, De Angeli & Tessier;
p. 196, Text-fig. 12, PI. 11 (figs. 1-2)
1994 - Ranilia punctulata Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 176, PI. 4 (fig. 3)
2001 - Ranilia punctulata Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 22, Text-figs. 14(1 a-b), 15
38
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Holotype: MCZ 1120 (I.G. 21 1656).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) reported four speci-
mens (SV from 311 to 313; MCZ 1120) from “Lovara”
and “Boschetto” quarries of Chiampo (Vicenza); Beschin
et al. (1994) reported three specimens (MCZ 1256, 1415,
1442) from “Boschetto” quarry of Nogarole Vicentino
(Vicenza).
Ranina Lamarck, 1 80 1
Type species: Ranina ranina (Linnaeus, 1758)
(= R. serrata Lamarck, 1801; R. dentata Latreille,
1802).
Stratigraphic range: Eocene - Recent.
Ranina bouilleana A. Milne Edwards, 1872
1872 - Ranina bouilleana A. Milne Edwards; p. 6, PI. 8 (figs. 2-
2 a-c)
1873 - Ranina bouilleana A. Milne Edwards in A. Milne Edwards;
p. 8, PI. 4 (fig. 55)
1 883 - Ranina bouilleana A. Milne Edwards in Bittner; p. 303
1 9 1 Oa - Ranina bouilleana A.. Milne Edwards in Fabiani; p. 20
191 Ob - Ranina ( Eteroranina ) bouilleana A. Milne Edwards in Fabia¬
ni; p. 91
1915 - Ranina bouilleana A. Milne Edwards in Fabiani; p. 285
1929 - Ranina (R.) bouilleana A. Milne Edwards in Glaessner; p. 361
2001 - Ranina bouilleana A. Milne Edwards in De Angeli & Beschin;
p. 20
Holotype: unknown.
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: Bittner (1883) reported one speciemen
from Castelli di Montecchio Maggiore (Vicenza) (cata¬
logne number unknown).
Note: this species is also reported from thè Oligocene
of France.
Ranina paìmea E. Sismonda, 1846
1846 - Ranina palmea E. Sismonda; p. 64, PI. 3 (figs. 3-4)
not 1847 - Ranina palmea E. Sismonda in Michelotti; p. 70 = Ranina
serrata
1 859 - Ranina palmea E. Sismonda in Reuss; p. 21
1872 - Ranina palmea E. Sismonda in A. Milne Edwards; p. 4
1895 - Ranina palmea E. Sismonda in Crema; p. 672, Text-
fig. 12 a-e
1 9 1 Ob - Ranina palmea E. Sismonda in Fabiani; p. 9
1 929 - Ranina palmea E. Sismonda in Glaessner; p. 362
Holotype: lost.
Stratigraphic range: Miocene.
Occurrence: Piemonte.
Material: E. Sismonda (1846) reported one specimen
from Torino, today lost; Crema (1895) reported some
specimens from Torino, Sciolze and Bardassano (Torino)
(repository and catalogne number unknown).
Ranina propinqua Ristori, 1891
1891 a - Ranina propinqua Ristori; p. 1 1, PI. 1 (figs. 4-7)
191 Ob — Ranina propinqua Ristori in Fabiani; p. 9
1 929 - Ranina propinqua Ristori in Glaessner; p. 363
Holotype: unknown.
Stratigraphic range: Pliocene.
Occurrence: Umbria.
Material: Ristori (189 la) reported one specimen from
Città della Pieve (Umbria) (MGPUR, catalogue number
unknown).
Ranina speciosa (Miinster, 1 840)
1840 - Hela speciosa Miinster; p. 24, PI. 2 (figs. 1-3)
1859 - Ranina speciosa (Miinster) in Reuss; p. 22
1875 - Ranina speciosa (Miinster) in Bittner; p. 70, PI. 1 (fig. 5 a-d)
1886 - Ranina speciosa (Miinster) in Noetling; p. 33
1 887 - Ranina speciosa (Miinster) in Ebert; p. 266, PI. 9 (fig. 1 )
1889 - Ranina speciosa (Miinster) in Ristori; p. 406, PI. 15
(figs. 9-13)
1903 - Ranina speciosa (Miinster) in Oppenheim; p. 196
191 Oa — Ranina speciosa (Miinster) in Fabiani; p. 20
191 Ob — Ranina speciosa (Miinster) in Fabiani; p. 8
1929 - Ranina speciosa (Miinster) in Glaessner; p. 363
1974 - Ranina speciosa (Miinster) in Mastrorilli; p. 4
1987 - Ranina speciosa (Miinster) in Allasinaz; p. 529, Text-fig. 9,
PI. 1 (figs. 11-16), PI. 2 (figs. 1-6), PI. 3 (figs. 1-8)
Holotype: unknown.
Stratigraphic range: Oligocene - Miocene.
Occurrence: Piemonte, Veneto, Liguria.
Material: Oppenheim (1903) and Bittner (1875)
reported this species from “Brocchi” quarry (Bas-
sano - Vicenza) and Monfumo (Treviso) respec-
tively (repository and catalogue number unknown);
Ristori (1889) described 13 specimens from Sassello
(Savona), housed in don Perrando’s collection today
in thè “DIP.TE.RIS” (we identified two originai speci¬
mens: 239 1/Sa-II-S 188 (ex Sa-II-S183) and 2392/Sa-
II-S189 (ex Sa-II-S 1 83), figured in PI. 15, figs. 9, 11)
and in Michelotti’s collection (SGR, catalogue number
unknown); Allasinaz (1987) reported 30 specimens
from Ciglione, Torrente Aimone and Rio Volpina,
and Ponzone (Acqui) (PU; MCSN, catalogue number
unknown).
Note: this species is also reported from Osnabriick and
Cassel (Germany).
Raninoides H. Milne Edwards, 1837
Type species: Ranina laevis Latreille, 1825.
Stratigraphic range: Eocene - Recent.
Note: Schweitzer et al. (in press) considered Lae-
viranina Lòrenthey (in Lòrenthey & Beurlen, 1929) as
synonym with Raninoides H. Milne Edwards, 1837.
Raninoides budapestiniensis (Lòrenthey, 1897)
1897 -Ranina budapestiniensis Lòrenthey; p. 153, 166
1898 - Ranina budapestinensis Lòrenthey in Lòrenthey; p. 23, PI. 1
(fig- 2)
1929 - Laeviranina budapestinensis (Lòrenthey) in Lòrenthey & Beur¬
len; p. 107, PI. 4 (fig. 12)
1929 - Ranina ( Laeviranina ) budapestinensis Lòrenthey in Glaessner;
p. 364
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
39
1994 - Laeviranina budapestiniensis (Lòrenthey) in Beschin, Busulini,
De Angeli & Tessier; p. 172, PI. 3 (fig. 1 a-c)
2001 - Laeviranina budapestiniensis (Lòrenthey) in De Angeli &
Beschin; p. 16
Holotype: unknown.
Stratigraphic range: middle/upper Eocene ( Lutetian -
et Priabonian).
Occurrence: Veneto.
Material; Beschin et al. (1994) reported one speci¬
men (MCZ 1295) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Raninoides fabianii (Lòrenthey & Beurlen, 1929)
1929 - Laeviranina Fabianii Lòrenthey & Beurlen; p. 105, PI. 4
(fig. 10)
1929 - Ranina ( Laeviranina ) fabianii Lòrenthey in Glaessner;
p. 364
193 1 - Laeviranina fabianii Lòrenthey & Beurlen in Glaessner & With-
ers; p. 491
1935 - Ranina ( Laeviranina ) fabianii Lòrenthey & Beurlen in Roger;
p. 352, PI. 9 (fig. 5)
1969 - Laeviranina fabianii Lòrenthey & Beurlen in Glaessner;
p. 364
1969 - Laeviranina fabianii Lòrenthey & Beurlen in Via Boada;
p. 391
1982 - Laeviranina fabianii Lòrenthey & Beurlen in Forster & Mund-
I los; p. 156, PI. 1 (figs. 4-6)
1998 - Laeviranina fabianii Lòrenthey & Beurlen in Tucker; p. 35 1
2006 - Laeviranina fabianii Lòrenthey & Beurlen in Beschin,
De Angeli, Checchi & Mietto; p. 103, PI. 2 (figs. 2-3)
i
Holotype: unknown.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (2006) reported six
specimens (MCZ from 2433 to 2438) from Val
Segato, Grotta della Poscola, and Buso della Rana
(Vicenza).
Note: this species is also reported from tha middle/
upper Eocene (Lutetian - Priabonian) of Hungary and
Germany.
Raninoides cfr. R. fabianii (Lòrenthey & Beurlen, 1929)
1998 - Laeviranina cfr. fabianii Lòrenthey & Beurlen in Beschin,
Busulini, De Angeli, Tessier & Ungaro; p. 1 8, Text-figs. 7, 9 (2)
2000 - Laeviranina cfr. fabianii Lòrenthey in Beschin, De Angeli &
Alberti; p. 15
2001 - Laeviranina cfr. fabianii Lòrenthey & Beurlen in De Angeli &
Beschin; p. 16
r
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1998) reported one speci¬
men (MCZ 1541) from “Rossi” quarry of Monte di Malo
(Vicenza).
Raninoides notopoides (Bittner, 1883)
1883 - Ranina notopoides Bittner; p. 304, PI. 1 (fig. 3)
191 Ob - Ranina ( Eteroranina ) notopoides Bittner in Fabiani;
p. 91
1915 - Ranina notopoides Bittner in Fabiani; p. 284
1929 - Ranina {R.) notopoides Bittner in Glaessner; p. 362
1969 - Laeviranina notopoides (Bittner) in Via Boada; p. 124, 388
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1883) reported one specimen from
Monte Masua of Negrar (Verona) (repository and cata-
logue number unknown).
Raninoides pulchra (Beschin, Busulini, De Angeli &
Tessier, 1988)
1988 - Laeviranina pulchra Beschin, Busulini, De Angeli & Tessier;
p. 171, Text-fig. 5 (2), PI. 4 (figs. 1-3)
1997 - Laeviranina pulchra Beschin, Busulini, De Angeli & Tessier in
Vicariotto; p. 29, Text-fig. 1 a-b
2001 - Laeviranina pulchra Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 16, Text-fig. 10 (6)
2004 - Laeviranina pulchra Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 113
Holotype: MCZ 1126 (I.G. 211662).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1988) ) reported three speci¬
mens (SV 250; MCZ 1126, 1129) from “Main” quarry
of Arzignano (Vicenza); Vicariotto (1997) reported one
specimen (MCZ 1576) from “Boschetto” quarry of Noga¬
role Vicentino (Vicenza).
Tribolocephalus Ristori, 1886
Type species: Tribolocephalus laevis Ristori, 1886.
Stratigraphic range: Pliocene.
Tribolocephalus laevis Ristori, 1886
1886 - Tribolocephalus laevis Ristori; p. 128, PI. 2 (fig. 19)
1927 - Tribolocephalus laevis Ristori in Van Straelen; p. 85
1929 - Tribolocephalus laevis Ristori in Glaessner; p. 388
1969 - Tribolocephalus laevis Ristori in Glaessner; p. 502
1981 - Tribolocephalus laevis Ristori in Delle Cave; p. 45
Holotype: IGF616E.
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (1886) reported one specimen (IGF
616E) from Orciano (Pisa).
Section Eubrachyura de Saint Laurent, 1980
Subsection Heterotremata Guinot, 1977
Superfamily Dorippoidea MacLeay, 1838
Family Dorippidae MacLeay, 1838
Medorippe Manning & Holthuis, 1 98 1
Type species: Dorippe lanata Linnaeus, 1767.
Stratigraphic range: Miocene - Recent.
40
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Medorippe ampia Garassino, De Angeli, Gallo & Pasini,
2004
2004 - Medorippe ampia Garassino, De Angeli, Gallo & Pasini;
p. 260, Text-figs. 5-6 a-b
Holotype: PU 41149.
Stratigraphic range: upper Miocene (Messinian).
Occurrence: Piemonte.
Material: Garassino et al. (2004) reported two speci-
mens (PU 4 1 1 49, 4 1 1 97) from Cocconato (Asti).
Medorippe lanata (Linnaeus, 1767)
1767 - Dorippe lanata Linnaeus; p. 1044
1863 - Dorippe lanata Linnaeus in Heller; p. 138, PI. 4 (fig. 9)
1914 - Dorippe lanata Linnaeus in M. Gemmellaro; p. 78, PI. 1
(fig- 5)
1929 - Dorippe lanata Linnaeus in Glaessner; p. 137
2004 - Medorippe lanata (Linnaeus) in Garassino, De Angeli, Gallo &
Pasini; p. 262
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported some
specimens, housed in Marchese di Monterosato’s collec-
tion (repository and catalogue number unknown).
tFamily Orithopsidae Schweitzer, Feldmann, Fam,
Hessin, Hetrick, Nyborg & Ross, 2003
Cherpiocarcinus Marangon & De Angeli, 1997
Type species: Cherpiocarcinus rostratus Marangon &
De Angeli, 1997.
Stratigraphic range: lower Oligocene (Rupelian).
Cherpiocarcinus rostratus Marangon & De Angeli,
1997
1997 - Cherpiocarcinus rostratus Marangon & De Angeli; p. 102,
Text-fig. 2, PI. 1 (figs. 1-2)
2003a - Cherpiocarcinus rostratus Marangon & De Angeli in
De Angeli & Marangon; p. 1 03, Text-fig. 1 (1)
2003 - Cherpiocarcinus rostratus Marangon & De Angeli in Sch¬
weitzer, Feldmann, Fam, Hessin, Hetrick, Nyborg & Ross; p. 39,
PI. 13 (fig. 1)
Holotype: MCZ 1548 (I.G. 284620).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte.
Material: Marangon & De Angeli (1997) reported
two specimens (MCZ 1548, 1549) from Case Cherpione
(Alessandria).
Superfamily Calappoidea A. Milne Edwards, 1837
Family Calappidae A. Milne Edwards, 1837
Bittnerilia De Angeli & Garassino, 2003
Type species: Lambrus eocaenus Bittner, 1883.
Stratigraphic range: middle Eocene (Lutetian).
Bittnerilia dentata Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 - Bittnerilia dentata Beschin, De Angeli, Checchi & Zaran¬
tonello; p. 17, Text-fig. 11, PI. 3 (fig. 7 a-b)
Holotype: MCZ 2298 (I.G. 296517).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported two speci¬
mens (MCZ 2298, 2319) from Grola di Comedo Vicen¬
tino (Vicenza).
Bittnerilia eocaena (Bittner, 1883)
1883 - Lambrus eocaenus Bittner; p. 309, PI. 1 (fig. 7 a-c)
191 Oa - Lambrus eocaenus Bittner in Fabiani; p. 28
1915 - Lambrus eocaenus Bittner in Fabiani; p. 284
1983 - Parthenope eocaena (Bittner) in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 62, PI. 2 (fig. 3 a-b)
1994 - Parthenope eocaena (Bittner) in Beschin, Busulini, De Angeli
& Tessier; p. 181, PI. 6 (fig. 1 a-b)
2001 - Parthenope eocaena (Bittner) in De Angeli & Beschin; p. 27
2003 - Bittnerilia eocaena (Bittner) in De Angeli & Garassino; p. 17,
Text-figs. 2-4
2004 - Bittnerilia eocaena (Bittner) in Beschin, Busulini, De Angeli &
Tessier; p. 1 15
2005 - Bittnerilia eocaena (Bittner) in Beschin, De Angeli, Checchi &
Zarantonello; p. 18
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1883) reported one specimen from
S. Giovanni Ilarione (Verona) (repository and catalogue
number unknown); Busulini et al. (1983) reported one
specimen (*18) from “Main” quarry of Arzignano
(Vicenza); Beschin et al. (1994) reported three speci¬
mens (MCZ 1289, 1290, 1293) from “Boschetto” quarry
of Nogarole Vicentino (Vicenza); De Angeli & Ga¬
rassino (2003) reported one specimen (MCZ 2387) from
“Albanello” quarry of Nogarole Vicentino (Vicenza).
Calappa Weber, 1795
Type species: Cancer granulatus Linnaeus, 1758.
Stratigraphic range: ?middle Eocene - Recent.
Calappa granulata (Linnaeus, 1758)
1758 - Cancer granulatus Linnaeus; p. 627
1767 - Cancer granulatus Linnaeus; p. 533
1798 - Calappa granulata (Linnaeus) in Fabricius; p. 346
1 8 1 6 - Calappa granulata (Linnaeus) in Risso; p. 1 8
1825 - Calappa granulata (Linnaeus) in Desmarest; p. 109, PI. 10
(fig- 1)
1828 - Calappa granulala (Linnaeus) in Roux; PI. 2 (fig. 13), PI. 16
(figs. 1-7)
1 861 - Calappa granulata (Linnaeus) in A. Milne Edwards; p. 88
1863 - Calappa granulala (Linnaeus) in Heller; p. 130, PI. 4 (fig. 3)
1914 - Calappa granulata (Linnaeus) in M. Gemmellaro; p. 80, PI. 1
(figs. 9-10)
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DEC APODA FROM ITALY
41
1918 - Calappa granulata (Linnaeus) in Pesta; Text-fig. 97
1929 - Calappa granulata (Linnaeus) in Glaessner; p. 70
1936 - Calappa granulala (Linnaeus) in Nobre; p. 81, PI. 28
(figs. 73-74)
1940 - Calappa granulata (Linnaeus) in Bouvier; p. 203, Text-fig. 203,
PI. 7 (fig. 1)
1946 - Calappa granulata (Linnaeus) in Zariquiey Alvarez; p. 143,
Text-fig. 163
1965 - Calappa granulata (Linnaeus) in Forest; p. 362
1968 - Calappa granulata (Linnaeus) in Zariquiey Alvarez; p. 315,
Text-figs. 1 05 c, 1 07 a
1992 - Calappa granulata (Linnaeus) in Falciai & Minervini; p. 181,
PI. 12 (fig. 4)
2004a - Calappa granulata (Linnaeus) in Garassino & De Angeli;
p. 38, Text-fig. 4(1-3)
2004 - Calappa granulata (Linnaeus) in Garassino, De Angeli, Gallo &
Pasini; p. 264, Text-fig. 7 a-c
Holotype: lost.
Stratigraphic range: Pliocene - Pleistocene.
Occurrence: Piemonte, Emilia Romagna, Sicilia.
Material; M. Gemmellaro (1914) reported two
specimens from Monte Pellegrino and Altavilla
(Palermo), thè first housed in Marchese di Montero-
sato’s collection (repository unknown) and thè second
in MGUP (catalogue number unknown); Garassino
& De Angeli (2004a) reported three specimens (MG
from 0609 to 0611) from Fiume Arda (Piacenza);
Garassino et al. (2004) reported 14 specimens (PU
from 41150 to 41163) from Candelo, Masserano and
Cossato (Biella).
Calappa sp.
1 89 la— Calappa sp. in Ristori; p. 9, PI. 1 (figs. 10, 15)
1 89 1 b — Calappa sp. in Ristori; p. 21
1929 - Calappa sp. in Glaessner; p. 72
Stratigraphic range; Pliocene.
Occurrence; Toscana, Lazio.
Material: Ristori (189 la) reported two specimens
from Orciano (Pisa), today lost; Ristori ( 1 89 1 b) reported
many specimens from Monte Mario (Roma), belonging
to Zuccari’s collection (repository and catalogue number
unknown).
Calappa sp.
1895 - Calappa sp. in Crema; p. 673, Text-fig. 13
1 929 - Calappa sp. in Glaessner; p. 7 1
Stratigraphic range: Miocene - Pliocene.
Occurrence: Piemonte.
Material: Crema (1895) reported some specimens
from Torino, Sciolze, Baldissero (Torino), and Pino
d’Asti (Asti), today lost.
Calappa sp.
1 896 - Calappa sp. in Ristori; p. 507, PI. 12 (fig. 1 1 )
1909 - Calappa sp. in Lòrenthey; p. 235, PI. 2 (fig. 7)
1929 - Calappa sp. in Glaessner; p. 72
1950 - Calappa sp. in Comaschi Caria; p. 326
1956 - Calappa sp. in Comaschi Caria; p. 288
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Ristori (1896) reported one specimen from
S. Guglielmo (Cagliari), today lost; Lòrenthey (1909)
reported some specimens from Cadreas (Sassari) and
Monte S. Michele (Cagliari), belonging to Lovisato’ col¬
lection (repository and catalogue number unknown).
Calappa sp.
1 909 - Calappa sp. in Lòrenthey; p. 236
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported some speci¬
mens from Nurri, Monte S. Michele, Bonorva and S.
Avendrace (Cagliari) (repository and catalogue number
unknown).
Calappa sp.
1987 - Calappa sp. in Allasinaz; p. 521, Text-fig. 4, PI. 1 (fig. 1)
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte.
Material: Allasinaz (1987) reported one specimen
(MCSN 256) from Fontanino di Rio Caramagna (Cas-
sinelle).
Calappa ? sp.
1907 - Calappa ? sp. in Lòrenthey; p. 82, PI. 4 (fig. 8)
1909 - Calappa sp. in Lòrenthey; p. 236, PI. 2 (fig. 8)
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported some speci¬
mens from S. Avendrace (Cagliari), housed in Lovi-
sato’s collection (repository and catalogue number
unknown).
Calappa spec.
1875 - Calappa spec. in Bittner; p. 74, PI. 1 (fig. 7 a-b)
1929 - Calappa sp. Bittner in Glaessner; p. 72
2001 - Calappa spec. Bittner in De Angeli & Beschin; p. 22
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875) reported one specimen from
S. Giovanni Barione (Verona) (repository and catalogue
number unknown).
Calappilia A. Milne Edwards, 1873
Type species: Calappilia verrucosa A. Milne
Edwards, 1873.
Stratigraphic range: Eocene - Recent.
Calappilia dacica Bittner, 1893
1893 - Calappilia dacica Bittner; p. 16, PI. 2 (fig. 1)
1898 - Calappilia dacica Bittner in Lòrenthey; p. 30, PI. 1 (fig. 5)
1929 - Calappilia dacica Bittner in Glaessner; p. 73
42
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1929 - Calappilia dacica Bittner in Lòrenthey & Beurlen; p. 127,
PI. 6 (figs. 4-6)
1982 - Calappilia dacica Bittner in Busulini, Tessier & Visentin; p. 77
2001 - Calappilia dacica Bittner in De Angeli & Beschin; p. 22
2004 - Calappilia dacica Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 1 15
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1982) reported two speci-
mens from “Main” quarry of Arzignano (Vicenza) (repos-
itory and catalogue number unknown).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Calappilia gemmata Beschin, Busulini, De Angeli &
Tessier, 1994
1994 - Calappilia gemmata Beschin Busulini, De Angeli & Tessier;
p. 167, Text-fig. 3, PI. 2 (figs. 1-2)
2001 - Calappilia gemmata Beschin Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 22
Holotype: MCZ 1427 (I.G. 284616).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported four speci-
mens (MCZ from 1425 to 1428) from “Boschetto” quarry
of Nogarole Vicentino (Vicenza).
Calappilia incisa Bittner, 1886
1886 - Calappilia incisa Bittner; p. 48, Text-fig. 3 a-c
191 Oa — Calappilia incisa Bittner in Fabiani; p. 6
1915 - Calappilia incisa Bittner in Fabiani; p. 284
1929 - Calappilia incisa Bittner in Glaessner; p. 73
1982 - Calappilia cfr. incisa Bittner in Busulini, Tessier & Visentin;
p. 77
1994 - Calappilia incisa Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 168, PI. 1 (figs. 5-6)
2001 - Calappilia incisa Bittner in De Angeli & Beschin; p. 22
2004 - Calappilia cfr. incisa Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 1 15
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1886) reported one specimen from
“Scole” quarry (Verona) (repository and catalogue number
unknown); Busulini et al. (1982) reported two specimens
from “Main” quarry of Arzignano (Vicenza) (repository
and catalogue number unknown); Beschin et al. (1994)
reported two specimens (MCZ 1259, 1446) from “Bos¬
chetto” quarry of Nogarole Vicentino (Vicenza).
Calappilia mainii Allasinaz, 1 987
1987 - Calappilia mainii Allasinaz; p. 523, Text-fig. 5, PI. 1
(figs. 2-6)
Holoty pe: MCSN 158 M.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte.
Material: Allasinaz (1987) reported five specimens
(MCSN 158 M, la, 189, 289, 162 M) from Ciglione ! :
(Alessandria).
Calappilia subovata Beschin, Busulini, De Angeli &
Tessier, 2002
2002 - Calappilia subovata Beschin, Busulini, De Angeli & Tessier;
p. 13, Text-fig. 8, PI. 2 (fig. 4)
2004 - Calappilia subovata Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 15
Holotype: MCZ 2269 (I.G. 296397).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported one speci¬
men (MCZ 2269) from “Main” quarry of Arzignano
(Vicenza).
Calappilia verrucosa A. Milne Edwards, 1873
1873 - Calappilia verrucosa A. Milne Edwards; p. 8, PI. 4 (figs. 3,
3 a)
2003a - Calappilia verrucosa A. Milne Edwards in De Angeli &
Marangon; p. 101, Text-fig. 1 (2)
2003b - Calappilia verrucosa A. Milne Edwards in De Angeli &
Marangon; p. 190, Text-figs. 3-5
Holotype: MNHN R03771 (cast).
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Piemonte.
Material: De Angeli & Marangon (2003 a, b) reported
two specimens (MCZ 2388, 2389) from Case Cherpione
(Alessandria).
Note: this species is also reported from thè Oligocene
of France.
Calappilia vicetina Fabiani, 1910
1910a - Calappilia vicetina Fabiani; p. 4, 21, PI. 1 (fig. 1 a-c)
1915 - Calappilia vicetina Fabiani in Fabiani; p. 285
1929 - Calappilia vicentina Fabiani in Glaessner; p. 74
1987 - Calappilia vicetina Fabiani in Allasinaz; p. 525, Text-fig. 6,
PI. 1 (fig- 7)
2001 - Calappilia vicetina Fabiani in De Angeli & Beschin; p. 23
Holotype: MGPD 23999.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte, Veneto.
Material: Fabiani (191 Oa) reported two specimens
(MGPD 23999, same catalogue number for thè two speci¬
mens) from Case Maraschin di Nanto and Case Soghe
(Monti Berici); Allasinaz (1987) reported two specimens
(MCSN 236; PU 10) from Ponzone (Acqui).
Calappilia sp.
1997 - Calappa heberti Brocchi in Vicariotto; p. 28, Text-fig. 2 a-d
2000 - Calappilia sp. in De Angeli & Franchi; p. 20
2001 - Calappilia sp. in De Angeli & Beschin; p. 23
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Vicariotto (1997) reported one specimen
CATALOGANO BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
43
' (MCZ 1 575) from “Boschetto” quarry of Nogarole Vicen-
ie tino (Vicenza).
Mursiopsis Ristori, 1889
Type species: Mursiopsis pustu/osus Ristori, 1889.
Stratigraphic range: lower Oligocene (Rupelian).
r’ Il
Mursiopsis pustulosus Ristori, 1889
n '
1889 - Mursiopsis pustulosus Ristori; p. 405, PI. 15 (figs. 6-8)
1929 - Mursiopsis pustulosus Ristori in Glaessner; p. 261
1969 - Mursiopsis pustulosus Ristori in Glaessner; p. 495
1974 - Mursiopsis pustulosus Ristori in Mastrorilli; p. 4
1987 - Mursiopsis pustulosus Ristori in Allasinaz; p. 527, Text-fig. 7,
PI. 1 (figs. 8-9)
Holotype: lost.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte, Liguria.
Material: Ristori (1889) reported one specimen from
Sassello (Savona), housed in don Perrando’s collection,
today lost; Allasinaz (1987) reported three specimens
(MCSN 361, 223, 101M), from Ciglione and Ponzone
(Acqui).
Mursiopsis sp.
1909 - Mursiopsis sp. in Lòrenthey; p. 236
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported one speci¬
men from Monte della Pace (Cagliari), housed in
Lovisato’s collection (repository and catalogue number
unknown).
Family Hepatidae Stimpson, 1871
Hepatiscus Bittner, 1875
Type species: Hepatiscus neumayri Bittner, 1875.
Stratigraphic range: Eocene.
? Hepatiscus distefanoi Checchia-Rispoli, 1905
1905 - Hepatiscus distefanoi Checchia-Rispoli; p. 323, PI. 1 (figs. 9,
11-13)
1929 - Hepatiscus distefanoi Checchia-Rispoli in Glaessner; p. 209
1969 - Hepatiscus? distefanoi Checchia-Rispoli in Via Boada;
p. 384
Holotype: MGUP.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1905) reported one
specimen from Pachino (Siracusa) (MGUP, catalogue
number unknown).
Note: Via Boada (1969) ascribed this species to thè
family Leucosiidae.
Hepatiscus minimus Beschin, Busulini, De Angeli &
Tessier, 1994
1994 - Hepatiscus minimus Beschin, Busulini, De Angeli & Tessier;
p. 169, Text-fig. 4, PI. 2 (figs. 4-5)
1999 - Hepatiscus minimus Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 13, Text-fig. 2 (5), PI. 2 (fig. 6)
2001 - Hepatiscus minimus Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 23
Holotype: MCZ 1429 (I.G. 284490).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported six speci¬
mens (MCZ 1429, 1439, 1344, 1456, 1458, 1459) from
“Boschetto” quarry of Nogarole Vicentino (Vicenza).
Hepatiscus neumayri Bittner, 1875
1875 - Hepatiscus neumayri Bittner; p. 75, PI. 1 (fig. 8)
1883 - Hepatiscus neumayri Bittner in Bittner; p. 312, PI. 1 (fig. 9)
1905 - Hepatiscus neumayri Bittner in Checchia-Rispoli; p. 324
191 Oa - Hepatiscus neumayri Bittner in Fabiani; p. 35, PI. 1 (fig. 9)
1915 - Hepatiscus Neumayri Bittner in Fabiani; p. 284
1933 - Hepatiscus neumayri Bittner in Di Salvo; p. 17
1982 - Hepatiscus neumayri Bittner in Busulini, Tessier & Visentin;
p. 81
1983 - Hepatiscus neumayri Bittner in Busulini, Tessier, Beschin,
Visentin, De Angeli & Rossi; PI. 3 (fig. 2)
1994 - Hepatiscus neumayri Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 169, PI. 2 (fig. 6)
1999 - Hepatiscus neumayri Bittner in De Angeli & Beschin; p. 14
Text-fig. 2 (2, 3, 4), PI. 1 (figs. 5 a-b, 6)
2001 - Hepatiscus neumayri Bittner in De Angeli & Beschin; p. 23,
Text-fig. 16
2004 - Hepatiscus neumayri Bittner in Beschin & De Angeli; p. 21
2004 - Hepatiscus neumayri Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 115
2005 - Hepatiscus neumayri Bittner in Beschin, De Angeli, Checchi &
Zarantonello; p. 18, PI. 3 (fig. 4)
Holotype: MNHB MB.A. 658.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto, Sicilia.
Material: Bittner (1875, 1883) reported one speci¬
men (MNHB MB.A. 658) from Ciupio (Verona); Fabiani
(1910a) reported two specimes (MGPD 11685, 11686)
from Ciupio and Buca del Prete (Verona); Di Salvo
(1933) reported one specimen from Ponte di Castronuovo
(Palermo) (MGUP, catalogue number unknown); Busu¬
lini et al. (1982) reported six specimens from “Main”
quarry of Arzignano (Vicenza), (repository and catalogue
number unknown); Beschin et al. (1994) reported one
specimen (MCZ 1211) from “Boschetto” quarry of Noga¬
role Vicentino (Vicenza); De Angeli & Beschin (1999)
reported 24 specimens (MCZ 1172, 1173, 1211, 1278,
1294, 1340, from 1625 to 1643) from Ciupio (Verona),
and “Main” quarry of Arzignano, “Boschetto” and
“Albanello” quarries of Nogarole Vicentino (Vicenza);
Beschin et al. (2005) reported 18 specimens (MCZ 2289,
2290, 2291, 2299, 2307, 2308, 2312, 2313, 2314, 2315,
2336, 2347, 2348, 2349, 2379; MCV 52, 04/12, 04/13)
from Grola di Comedo Vicentino (Vicenza).
Hepatiscus pulchellus Bittner, 1875
1875 - Hepatiscus pulchellus Bittner; p. 75, PI. 1 (figs. 9 a-c, 10)
44
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1905 - Hepatiscus pulchellus Bittner in Checchia-Rispoli; p. 324
191 Oa - Hepatiscus pulchellus Bittner in Fabiani; p. 8, 21, 29, PI. 1
(figs. 3-4)
1915 - Hepatiscus pulchellus Bittner in Fabiani; p. 284
1 929 - Hepatiscus pulchellus Bittner in Glaessner; p. 209
1983 - Hepatiscus pulchellus Bittner in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 64, PI. 3 (fig. 5)
1 994 - Hepatiscus pulchellus Bittner in Beschin, Busulini, De Angeli &
Tessier; p. 171, PI. 2 (fig. 3)
1999 - Hepatiscus pulchellus Bittner in De Angeli & Beschin; p. 16,
Text-fig. 2 (6), PI. 1 (figs. 3-4)
2001 - Hepatiscus pulchellus Bittner in De Angeli & Beschin; p. 23
2004 - Hepatiscus pulchellus Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 1 1 5
2005 - Hepatiscus pulchellus Bittner in Beschin, De Angeli, Checchi &
Zarantonello; p. 19, PI. 3 (fig. 6)
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875) reported two specimens
from Ciupio (Verona) (repository and catalogue number
unknown); Fabiani (1910a) reported two specimens
(MGPD 11770, 13808) from Ciupio (Verona); Busulini et
al. (1983) reported three specimens (*30, *17; MSNVE
10094) from “Main” quarry of Arzignano (Vicenza);
Beschin et al. (1994) reported one specimen (MCZ 1267)
from “Boschetto” quarry of Nogarole Vicentino (Vicen¬
za); De Angeli & Beschin (1999) reported five specimens
(MCZ 1267, from 1644 to 1647) from “Main” quarry
of Arzignano (Vicenza), “Boschetto” and “Albanello”
quarries of Nogarole Vicentino (Vicenza); Beschin et al.
(2005) reported four specimens (MCZ 2292, 2350, 2382;
MV 53) from Grola di Comedo Vicentino (Vicenza).
Note: this species is also reported from thè middle
Eocene (Lutetian) of N Africa.
Hepatiscus sp.
1982 - Hepatiscus sp. in Busulini, Tessier & Visentin; p. 81
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1982) reported three speci¬
mens from “Main” quarry of Arzignano (Vicenza) (SV,
catalogue number unknown).
Mainhepatiscus De Angeli & Beschin, 1999
Type species: Mainhepatiscus zannatoi De Angeli &
Beschin, 1999.
Stratigraphic range: middle Eocene (Lutetian).
Mainhepatiscus zannatoi De Angeli & Beschin, 1999
1 999 - Mainhepatiscus zannatoi De Angeli & Beschin; p. 1 8, Text-fig.
2(8), PI. 2 (figs. 1-2)
2001 - Mainhepatiscus zannatoi De Angeli & Beschin in De Angeli &
Beschin; p. 24, Text-fig. 17
2004 - Mainhepatiscus zannatoi De Angeli & Beschin in Beschin,
Busulini, De Angeli & Tessier; p. 1 1 5
Holotype: MCZ 1619 (I.G. 284507).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: De Angeli & Beschin (1999) reported three
specimens (MCZ 1182, 1619, 1620) from “Main” quarry
of Arzignano (Vicenza).
Osachila Stimpson, 1871
Type species: Osachila tuberosa Stimpson, 1871.
Stratigraphic range: Eocene - Recent.
Osachila berica De Angeli & Beschin, 1999
1999 - Osachila berica De Angeli & Beschin; p. 18, Text-fig. 2 (9),
PI. 2 (figs. 3-4)
2001 - Osachila berica De Angeli & Beschin in Beschin, De Angeli &
Checchi; p. 29
2001 - Osachila berica De Angeli & Beschin in De Angeli & Beschin;
p. 24
2004 - Osachila berica De Angeli & Beschin in Beschin & De Angeli;
P- 21
Holotype: MCZ 1622 (LG. 284510).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Beschin (1999) reported three
specimens (MCZ 1622, 1623, 1624) from Campolongo
(Vicenza).
Priabonella Beschin, De Angeli, Checchi & Mietto,
2006
Type species: Priabonella violata Beschin, De Angeli,
Checchi & Mietto 2006.
Stratigraphic range: Eocene.
Priabonella violata Beschin, De Angeli, Checchi &
Mietto 2006
1999 - Hepatiscus poverelli Via Boada in De Angeli & Beschin; p.13,
Text-fig. 2 (1), PI. 1 (figs. 1-2)
2006 - Priabonella violatii Beschin, De Angeli, Checchi & Mietto;
p. 104, Text-fig. 6, PI. 2 (figs. 4-6)
Holotype: MCZ 2040.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Beschin (1999) reported 11
specimens (MCZ from 1555 to 1561, from 1615 to 1618)
from “Alonte” quarry (Monti Berici, Vicenza), and two
specimens (MCZ 1555, 1556) from S. Bovo (Bassano del
Grappa) and Val Segato (Vicenza); Beschin et al. (2006)
reported two specimens (MCZ 2040; MGPD 28739) from
Rio Rana and Val Segato (Vicenza).
Pseudohepatiscus Blow & Manning, 1996
Type species: Pseudohepatiscus mari noi Blow &
Manning, 1996.
Stratigraphic range: Eocene.
Pseudohepatiscus silvanoi De Angeli & Beschin, 1 999
1999 - Pseudohepatiscus silvanoi De Angeli & Beschin; p. 20, Text-
fig. 2 (7), PI. 2 (fig. 5)
CATALOGANO BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
45
2001 - Pseudohepatiscus silvanoi De Angeli & Beschin in De Angeli
& Beschin; p. 24
Holotype: MCZ 1621 (I.G. 284509).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: De Angeli & Beschin (1999) reported one
specimen (MCZ 1621) from “Alvese” quarry (Vicenza).
Superfamily Leucosioidea Samouelle, 1819
Family Leucosiidae Samouelle, 1819
Ebalia Leach, 1817
Type species: Cancer tuberosus Pennant, 1777.
Stratigraphic range: Miocene - Recent.
Ebalia cranchii Leach, 1817
1817 - Ebalia cranchii Leach; PI. 25 (figs. 7-11)
1 846 - Ebalia cranchii Leach in Bell; p. 148
1855 - Ebalia cranchii Leach in Bell; p. 303
1 89 1 b - Ebalia cranchii var. romana Ristori; p. 22, PI. 1 (figs. 21-23),
nov. syn.
1 892a - Ebalia cranchii Leach in Ristori; p. 88
1914 - Ebalia cranchii var. romana Ristori in M. Gemmellaro; p. 78,
PI. 1 (figs. 3-6)
1929 - Ebalia cranchii var. romana Ristori in Glaessner; p. 144
1936 - Ebalia cranchii Leach in Nobre; p. 78, PI. 26 (fig. 64)
1940 - Ebalia cranchii Leach in Bouvier; p. 209, PI. 7 (figs. 7-10)
1946 - Ebalia cranchii var. romana Ristori in Maxia; p. 132, PI. 1
(fig- 1)
1956 - Ebalia cranchii Leach in Monod; p. 122, Text-figs. 145-146
1965 - Ebalia cranchii Leach in Forest; p. 364-371, Text-fig. 25 a-b,
PI. 2 (figs. 1-2)
1968 - Ebalia cranchii Leach in Zariquiey Alvarez; p. 329, Text-
figs. 108 a-b, 111 b, d, 111 A, a, 111 C, b
1981 - Ebalia cranchii var. romana Ristori in Delle Cave; p. 45
1992 - Ebalia cranchii Leach in Falciai & Minervini; p. 184
2004a - Ebalia cranchii Leach in Garassino & De Angeli; p. 39, Text-
figs. 3 (7, 8), 4 (4), 5, 6
Holotype: unknown.
Stratigraphic range: Pliocene - Pleistocene.
Occurrence: Emilia Romagna, Lazio, Sicilia.
Material: Ristori ( 1 89 1 b) reported some specimens
(IGF 991 E, syntype) from Farnesina and Monte Mario
(Roma), housed in Zuccari’s and Clerici’s collections,
today lost; M. Gemmellaro (1914) reported some speci¬
mens from Monte Pellegrino and Ficarazzi (Palermo),
housed in Marchese di Monterosato’s collection (reposi-
tory and catalogne number unknown) and in MGUP
(catalogue number unknown); Maxia (1946) reported one
specimen from Monte Mario (Roma), housed in Clerici’s
collection (MGPUR, catalogue number unknown); Ga¬
rassino & De Angeli (2004a) reported 14 specimens (MG
from 0604 to 0608, from 0622 to 0630, 0648) from Fiume
Arda (Piacenza).
Ebalia fuciriii Ristori, 1 892
1892a -Ebalia fucinii Ristori; p. 88, Text-fig. 2
1929 - Ebalia fucinii Ristori in Glaessner; p. 144
1981 - Ebalia fucinii Ristori in Delle Cave; p. 45
2004 - Ebalia fucinii Ristori in Garassino & De Angeli; p. 40
Holotype: lost.
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (1892a) reported one specimen from
Spicchio (Empoli), housed in Fucini’s collection, today
lost.
Ebalia lamarmorai Lòrenthey, 1909
1909 -Ebalia lamarmorai Lòrenthey; p. 232, PI. 1 (figs. 2-3 a-b)
Holotype: unknown.
Stratigraphic range: middle Miocene (Langhian).
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported two specimens
from Monte S. Michele (Cagliari), housed in Lovisato’s
collection (repository and catalogue number unknown).
Ebalia tuberosa Pennant, 1777
1777 - Cancer tuberosus Pennant; PI. 19 a (fig. 19)
1817 - Ebalia pennanti Leach; PI. 25 (figs. 1-6), nov. syn.
1 89 1 b - Ebalia pennantii Leach in Ristori; p. 21, PI. 1 (figs. 24-25)
1893 - Ebalia pennanti Leach in Ristori; p. 88
1897 - Ebalia pennanti Leach in Bell; p. 3
1914 - Ebalia pennanti Leach in M. Gemmellaro; p. 79, PI. 1
(figs. 7-8)
1918 - Ebalia tuberosa (Pennant) in Pesta; p. 297, Text-fig. 91
1921 - Ebalia pennanti Leach in Bell; p. 8
1929 - Ebalia pennanti Leach in Glaessner; p. 145
1940 - Ebalia tuberosa (Pennant) in Ferrer Galdiano; p. 76, Text-
figs. 3-6
1940 - Ebalia tuberosa (Pennant) in Bouvier; p. 207, 211, PI. 7
(figs. 21-25)
1946 - Ebalia tuberosa (Pennant) in Zariquiey Alvarez; p. 147, Text-
fig. 164 a, PI. 10 (b)
1 956 - Ebalia tuberosa (Pennant) in Monod; p. 1 24, Text-figs. 150-151
1965 - Ebalia tuberosa (Pennant) in Forest; p. 370
1968 - Ebalia tuberosa (Pennant) in Zariquiey Alvarez; p. 326, Text-
figs. 109 d, 110 a-c, 111 a
1992 - Ebalia tuberosa (Pennant) in Falciai & Minervini; p. 184, 185
2004a - Ebalia tuberosa (Pennant) in Garassino & De Angeli; p. 40
Holotype: unknown.
Stratigraphic range: Pliocene - Pleistocene.
Occurrence: Lazio, Sicilia.
Material: Ristori ( 1 89 1 b) reported some specimens
from Farnesina and Monte Mario (Roma), housed in Zuc¬
cari’s collection (MGPUR, catalogue number unknown);
M. Gemmellaro (1914) reported one specimen from Fica-
razzi (Palermo), housed in Marchese di Monterosato’s
collection (repository and catalogue number unknown).
Hepatìnulits Ristori, 1 886
Type species: Hepatinulus seguentiae Ristori, 1886.
Stratigraphic range: Miocene - Pliocene.
Hepatinulus lovisatoi Lòrenthey, 1909
1909 - Hepatinulus lovisatoi Lòrenthey; p. 230, PI. 1 (fig. 8)
46
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1929 -Hepatinulus lovisatoi Lòrenthey in Glaessner; p. 209
Holotype: unknown.
Stratigraphic range: lower Miocene (Burdigalian).
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported one
specimen from Cugia (Sassari), housed in Lovi-
sato’s collection (repository and catalogue number
unknown).
Hepatinulus seguentiae Ristori, 1 886
1886 - Hepatinulus seguentiae Ristori; p. 121, PI. 3 (figs. 6-7)
1929 - Hepatinulus seguentiae Ristori in Glaessner; p. 209
Holotype: unknown.
Stratigraphic range: Pliocene.
Occurrence: Sicilia.
Material: Ristori (1886) reported two specimens from
Scoppo and S. Filippo (Trapani) (repository and catalogue
number unknown).
Ilio Leach, 1817
Type species: Cancer nucleus Linnaeus, 1758.
Stratigraphic range: ?Eocene - Recent.
Ilia cfr. I. nucleus (Linnaeus, 1758)
1 89 lb - Ilia cfr. nucleus (Linnaeus) in Ristori; p. 23
1929 - Ilia cfr. nucleus (Linnaeus) in Glaessner; p. 225
Stratigraphic range: Pliocene.
Occurrence: Lazio.
Material: Ristori ( 1 89 1 b) reported three specimens
from Monte Mario (Roma) (Zuccari’s collection) (reposi¬
tory and catalogue number unknown).
Ilia pliocaenica Ristori, 1891
189 la - Ilia pliocaenica Ristori; p. 10, PI. 1 (figs. 8-9, 11-12, 14)
1 892a - Ilia pliocaenica Ristori in Ristori; p. 86
1929 - Ilia pliocaenica Ristori in Glaessner; p. 225
1981 - Ilia pliocaenica Ristori in Delle Cave; p. 45
2004a - Ilia pliocaenica Ristori in Garassino & De Angeli; p. 40, Text-
fig.4(5)
2004 - Ilia pliocaenica Ristori in Garassino, De Angeli, Gallo & Pasini;
p. 266, Text-fig. 8 a-b
Syntype: IGF 939E.
Stratigraphic range: Pliocene - Pleistocene.
Occurrence: Piemonte, Emilia Romagna, Toscana.
Material: Ristori (189 la) reported some speci¬
mens from Spicchio (Empoli) (IGF); Garassino & De
Angeli (2004a) reported five specimens (MG from
0636 to 0639, 0647) from Fiume Arda (Piacenza);
Garassino et al. (2004) reported six specimens (PU
from 41164 41169) from Masserano and Cossato
(Biella).
Nucilobus Morris & Collins, 1991
Type species: Nucilobus symmetricus Morris & Col¬
lins, 1991.
Stratigraphic range: upper Eocene - Neogene.
Nucilobus bericus De Angeli & Beschin, 2004
2004 - Nucilobus bericus De Angeli & Beschin; p. 120, Text-
figs. 2-3
Holotype: MCZ 2395 (I.G. 296613).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Beschin (2004) reported one
specimen (MCZ 2395) from Lonigo (Vicenza).
Palaeomyra A. Milne Edwards in E. Sismonda, 1861
Type species: Palaeomyra bispinosa A. Milne
Edwards in E. Sismonda, 1861.
Stratigraphic range: Oligocene - Miocene.
Palaeomyra bispinosa A. Milne Edwards in E.
Sismonda, 1861
1861 - Palaeomyra bispinosa A. Milne Edwards in E. Sismonda;
p. 16, Text-figs. 18-20
1969 - Palaeomyra bispinosa A. Milne Edwards in Glaessner;
p. R498
2004 - Palaeomyra bispinosa A. Milne Edwards in Garassino,
De Angeli, Gallo & Pasini; p. 267 , Text-figs. 9-10 a-b
Holotype: lost.
Stratigraphic range: Oligocene - Miocene.
Occurrence: Piemonte.
Material: E. Sismonda (1861) reported one specimen
from Torino, today lost; Garassino et al. (2004) reported
six specimens (PU from 41 170 to 41175) from Cocconato
(Asti) and Morbello (Alessandria).
Typilobus Stoliczka, 1871
Type species: Typilobus granulosus Stoliczka, 1871.
Stratigraphic range: Eocene - Miocene.
Typilobus semseyanus Lòrenthey, 1 897
1897 - Typilobus semseyanus Lòrenthey; p. 97
1898 - Typilobus semseyanus Lòrenthey in Lòrenthey; p. 27, PI. 1
(figs. 3-4)
1929 - Typilobus semseyanus Lòrenthey in Lòrenthey & Beurlen;
p. 125, PI. 4 (figs. 10-11)
1998 - Typilobus semseyanus Lòrenthey in Beschin, Busulini,
De Angeli & Ungaro; p. 1 7, Text-fig. 6 (4)
2000 - Typilobus semseyanus Lòrenthey in Beschin, De Angeli &
Alberti; p. 15
Holotype: unknown
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (1998) reported one speci¬
men (MCZ 1485) from “Rossi” quarry of Monte di Malo
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
CATALOG AND BTBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
47
Superfamily Majoidea Samouelle, 1819
Family Majidae Samouelle, 1819
Maja Lamarck, 1801
Type species: Cancer squinado Herbst, 1788.
Stratigraphic range: Miocene - Recent.
Maja miocaenica Lòrenthey, 1907
1907 -Maja miocaenica Lòrenthey; p. 83, PI. 3 (figs. 1, 10)
1909 - Maja miocaenica Lòrenthey in Lòrenthey; p. 237, PI. 1
(figs. 1 a-b, 10 a-f)
1929 - Maja miocaenica Lòrenthey in Glaessner; p. 247
1990 - Maja miocaenica Lòrenthey in Moissette & Miiller; p. 739,
PI. 1 (fig. 2)
Holotype: unknown.
Stratigraphic range: upper Miocene (Tortonian).
Occurrence: Sardegna.
Material: Lòrenthey (1907) reported two specimens
from Capo S. Marco (Oristano), housed in Lovisato’s col-
lection (repository and catalogue number unknown).
Note: this species is also reported from thè upper
Miocene of Oranie (Algérie) and Malta.
Maja squinado (Herbst, 1788)
1788 - Cancer squinado Herbst; p. 214, PI. 14 (figs. 84-85)
1 8 16 - Maja squinado (Herbst) in Risso; p. 44
1 8 1 7 - Maja squinado (Herbst) in Leach; p. 1 8 1 7, PI. 18 (figs. 1 -6)
1 825 - Maja squinado (Herbst) in Desmarest; p. 145
1 857 - Maja squinado (Herbst) in Meneghini; p. 557
1 861 - Maja squinado (Herbst) in A. Milne Edwards; p. 88
1863 - Maja squinado (Herbst) in Heller; p. 49, PI. 1 (figs. 17-24)
1871 - Maja squinado (Herbst) in Woodward; p. 130
1 855 - Maja squinado (Herbst) in Carus; p. 507
1 890 - Maja squinado (Herbst) in Reid; p. 28 1
1 89 1 b - Pseudocarcinusl Ristori; p. 20, PI. 1 (fig. 29), nov syn.
1897 - Maja squinado (Herbst) in Bell; p. 3
1903 - Maja squinado (Herbst) in Checchia-Rispoli; p. 488
1909 - Maja squinado (Herbst) in Lòrenthey; p. 238
191 1 - Maja squinado (Herbst) in Magri; p. 8
1914 - Maja squinado (Herbst) in M. Gemmellaro; p. 81, PI. 1
(figs. 11-12)
1914 - Maja squinado var. di-stefanoi M. Gemmellaro; p. 83, PI. 1
(figs. 13-14), nov. syn.
1918 - Maja squinado (Herbst) in Pesta; p. 361 , Text-fig. 1 16
1929 - Maja squinado (Herbst) in Glaessner; p. 247
1929 - Maja squinado (Herbst) in Santucci; p. 63
193 1 - Maja squinado (Herbst) in Nobre; p. 142, Text-fig. 82
1940 - Maja squinado (Herbst) in Bouvier; p. 321, Text-fig. 95
1946 - Maja squinado (Herbst) in Zariquiey Alvarez; p. 169, Text-
fig. 171
195 1 - Maja squinado (Herbst) in Capart; p. 98, Text-fig. 32
1956 - Maja squinado (Herbst) in Monod; p. 474, Text-figs. 638-643
1961 - Maja squinado (Herbst) in Luther & Fiedler; p. 117, 166
1966 - Maja squinado (Herbst) in Bauchau; p. 79
1968 - Maja squinado (Herbst) in Zariquiey Alvarez; p. 446, Text-
figs. 149 a, 150 g, h
1980 - Maja squinado (Herbst) in Ingle; p. 23, 45, 141
1981 - Maja squinado (Herbst) in Varola; p. 13, PI. 3 (fig. 1)
1992 - Maja squinado (Herbst) in Falciai & Minervini; p. 247, PI. 17
(fig- 2)
2004a - Maja squinado (Herbst) in Garassino & De Angeli; p. 40,
Text-fig. 7
Holotype: unknown.
Stratigraphic range: Pliocene - Pleistocene.
Occurrence: Emilia Romagna, Lazio, Puglia, Sicilia,
Sardegna.
Material: Meneghini (1857) reported some
specimens from Alghero (repository and catalogue
number unknown); Ristori (1891 b) reported three
specimens from Monte Mario and Farnesina (Roma)
(repository and catalogue number unknown); M.
Gemmellaro (1914) reported one specimen from
Monte Pellegrino (Palermo) (MGUP, catalogue
number unknown); Varola (1981) reported one speci¬
men from Rocca Vecchia (Lecce) (GNSL, catalogue
number unknown); Garassino & De Angeli (2004a)
reported one specimen (MG 0603) from Fiume Arda
(Piacenza).
Maja sp.
1981 - Maja sp. in Varola; p. 14, PI. 3 (figs. 2-4)
Stratigraphic range: Pliocene.
Occurrence: Puglia.
Material: Varola (1981) reported three specimens from
Leuca (Lecce) (GNSL, catalogue number unknown).
Micromaia Bittner, 1875
Type species: Micromaia tubercolata Bittner, 1875.
Stratigraphic range: middle Eocene - lower Oli¬
gocene.
Micromaia elegans Beschin, Busulini, De Angeli &
Tessier, 1985
1985 - Micromaia elegans Beschin, Busulini, De Angeli & Tessier;
p. 102, Text-figs. 3 (7), 4 (4), PI. 2 (figs. 1 a-c, 2)
2001 - Micromaia elegans Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 26, Text-fig. 18 (6)
2002 - Micromaia elegans Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 14, Text-fig. 9, PI. 3
(fig- 3)
2004 - Micromaia elegans Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 15
Holotype: MSNVE 10482.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1985) reported three speci¬
mens (*197, *198; MSNVE 10482) from “Main” quarry
of Arzignano (Vicenza); Beschin et al. (2002) reported
one specimen (MCZ 2270) from “Main” quarry of Arzi¬
gnano (Vicenza).
Micromaia mainensis Beschin, Busulini, De Angeli &
Tessier, 1985
\9H5- Micromaia mainensis Beschin, Busulini, De Angeli & Tessier;
p. 104, Text-figs. 3 (4), 4 (2), PI. 3 (figs. 1-3)
2001 - Micromaia mainensis Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 26, Text-fig. 18 (4)
48
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
2004 - Micromaia mainensis Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 15
2005 - Micromaia mainensis Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli, Checchi & Zarantonello; p. 20, PI. 4 (fig. 2)
Holotype: MSNVE 10843.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1985) reported 19 speci¬
men (*98, *101, *102, * 104, *105, *106, *116, *117,
*118, *119, *191, *192, *193, *194, *195, *196, *214,
*264; MSNVE 10483) from “Main” quarry of Arzignano
(Vicenza); Beschin et al. (2005) reported eight specimens
(MCZ from 2294 to 2297, 2337, 2386; MV 49, 04/16)
from Grola di Comedo Vicentino (Vicenza).
Micromaia margaritata Fabiani, 1910
1910a - Micromaja margaritata Fabiani; p. 10, PI. 2 (fig. 2)
1915 - Micromaya margaritata Fabiani in Fabiani; p. 284
1929 - Micromaia margaritata Fabiani in Glaessner; p. 256
1982 - Micromaia margaritata Fabiani in Busulini, Tessier & Visentin;
p. 79
1985 - Micromaia margaritata Fabiani in Beschin, Busulini, De Angeli
& Tessier; p. 106, Text-fig. 3 (3), PI. 1 (figs. 3-4)
1 994 - Micromaia margaritata Fabiani in Beschin, Busulini, De Angeli
& Tessier; p. 179, PI. 5 (fig. 3)
2001 - Micromaia margaritata Fabiani in De Angeli & Beschin; p. 26,
Text-fig. 18 (3)
2004 - Micromaia margaritata Fabiani in Beschin, Busulini, De Angeli
& Tessier; p. 1 15
2005 - Micromaia margaritata Fabiani in Beschin, De Angeli, Checchi
& Zarantonello; p. 21, PI. 4 (fig. 3)
Holotype: MGPD 11687.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Vicenza.
Material: Fabiani (191 Oa) reported one specimen
(MGPD 11687) from Ciupio (Verona); Busulini et al.
(1982) reported two specimens from “Main” quarry of
Arzignano (Vicenza), (repository and catalogue number
unknown); Beschin et al. (1985) reported two specimens
(*27, *265) from “Main” quarry of Arzignano (Vicenza);
Beschin et al. (1994) reported three specimens (MCZ
1208, 1260, 1449) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza); Beschin et al. (2005) reported one
specimen (MCZ 2362) from Grola di Comedo Vicentino
(Vicenza).
Note: this species is also reported from thè Eocene of
Spain.
Micromaia meneguzzoi Beschin, Busulini, De Angeli &
Tessier, 1985
1985 - Micromaia meneguzzoi Beschin, Busulini, De Angeli &
Tessier; p. 107, Text-figs. 3 (5), 4 (3), PI. 4 (fig. 1 a-e)
2001 - Micromaia meneguzzoi Beschin, Busulini, De Angeli & Tessier
in De Angeli & Beschin; p. 26, Text-fig. 18 (5)
2004 - Micromaia meneguzzoi Beschin, Busulini, De Angeli & Tessier
in Beschin, Busulini, De Angeli & Tessier; p. 1 15
Holotype: MSNVE 10484.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1985) reported one speci¬
men (MSNVE 10484) from “Main” quarry of Arzignano
(Vicenza).
Micromaia priabonensis Oppenheim, 1901
1901 - Micromaja (?) priabonensis Oppenheim; p. 282, PI. 7
(figs. 13-1 3b)
1915 - Micromava priabonensis Oppenheim in Fabiani; p. 285
1929 - Micromaia (?) priabonensis Oppenheim in Glaessner; p. 257
1962 - Micromaja (?) priabonensis Oppenheim in Piccoli & Mocellin;
p. 86
1985 - Micromaia priabonensis Oppenheim in Beschin, Busulini,
De Angeli & Tessier; p. 109, Text-figs. 3-6, PI. 4 (figs. 2-3)
2001 - Micromaia priabonensis Oppenheim in De Angeli & Beschin;
p. 26
2006 - Micromaia priabonensis Oppenheim in Beschin, De Angeli,
Checchi & Mietto; p. 106, PI. 2 (fig. 7)
Holotype: unknown
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Oppenheim (1901) reported this species
from Priabona (Vicenza) (repository and catalogue
number unknown); Beschin et al. (1985) reported nine
specimens (*121, *213, *215, *221, *222, *223, *224,
*225, *266) from Priabona and Nanto (Vicenza); Beschin
et al. (2006) reported ten specimens (MCZ 1160, 1161,
1162, 1238, 1327, 2441, 2442, 2443, 2454, 2455) from
Buso della Rana (Vicenza).
Micromaia tuberculata Bittner, 1875
1875 - Micromaia tuberculata Bittner; p. 76, PI. 2 (fig. 2)
1883 - Micromaja tuberculata Bittner in Bittner; p. 308, PI. 1 (fig. 6)
1898 - Micromaia tuberculata Bittner in Lòrenthey; p. 31, PI. 1
(figs. 2-3)
1899 - Micromaia tuberculata Bittner in Oppenheim; p. 57
1909 - Micromaia tuberculata Bittner in Lòrenthey; p. 123
191 Oa - Micromaja tuberculata Bittner in Fabiani; p. 12, 21, PI. 2
(figs. 3-4)
1915 - Micromaya tuberculata Bittner in Fabiani; p. 284
1925 - Micromaia tuberculata Bittner in Schlosser; p. 143
1929 - Micromaia tuberculata Bittner in Glaessner; p. 257
1929 - Micromaia tuberculata Bittner in Lòrenthey & Beurlen; p. 141,
PI. 7 (figs. 4-5)
1969 - Micromaia tuberculata Bittner in Via Boada; p. 162, PI. 11
(figs. 1-4), PI. 12 (fig. 1)
1981 - Micromaia tuberculata Bittner in Quayle & Collins; p. 744,
PI. 104 (fig. 15)
1982 - Micromaia tuberculata Bittner in Busulini, Tessier, Visentin;
p. 79
1985 - Micromaia tuberculata Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 101, Text-figs. 3 (1), 4 (1), PI. 1, (figs. 1-2)
1994 - Micromaia tuberculata Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 178, PI. 5 (figs. 2-4)
2001 - Micromaia tuberculata Bittner in De Angeli & Beschin; p. 24,
Text-figs. 18(1), 19
2004 - Micromaia tuberculata Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 1 15
2005 - Micromaia tuberculata Bittner in Beschin, De Angeli, Checchi
& Zarantonello; p. 20, PI. 4 (fig. 1)
Holotype: unknown.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
49
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875, 1883) and Fabiani (191 Oa)
reported this species from S. Giovanni Barione (Verona)
(repository and catalogue number unknown); Busulini et
al. (1982) reported this species from “Main” quarry of
Arzignano (Vicenza) (repository and catalogue number
unknown); Beschin et al. (1985) reported 26 specimens
{*19, *81, *82, *83, *86, *87, *88, *89, *90, *92, *93,
*199, *200, *201, *202, *203, *204, *205, *206, *208,
*209, *210, *211, *220, *262, *263) from “Main”
quarry of Arzignano (Vicenza), “Albanello” quarry of
Nogarole Vicentino (Vicenza), Ciupio and Croce Grande
(Verona); Beschin et al. (1994) reported four specimens
(MCZ 1206, 1207, 1277, 1305) from “Boschetto” quarry
of Nogarole Vicentino (Vicenza); Beschin et al. (2005)
reported two specimens (MCZ 2335, 2346) from Grola di
Comedo Vicentino (Vicenza).
Note: this species is also reported from thè Eocene of
Hungary, Spain, Great Britain, and Egypt.
Micromaia sp.
1985 - Micromaia sp. in Beschin, Busulini, De Angeli & Tessier;
p. 110, Text-fig. 3 (2), PI. 2 (fig. 3 a-b)
2001 - Micromaia sp. in Beschin, De Angeli & Checchi; p. 29
2001 - Micromaia sp. in De Angeli & Beschin; p. 26
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: Beschin et al. (1985) reported one specimen
(MSNVE 10485) from Valmarana (Vicenza).
Periacanthus Bittner, 1875
Type species: Periacanthus horridus Bittner, 1875.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Periacanthus dallagoi Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 - Periacanthus dallagoi Beschin, De Angeli, Checchi & Zaran¬
tonello; p. 19, Text-fig. 12, PI. 3 (fig. 8)
Holotype: MCZ 2293 (I.G. 296512).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported three speci¬
mens (MCZ 2293, 2320, 2343) from Grola di Comedo
Vicentino (Vicenza).
Periacanthus horridus Bittner, 1875
1875 - Periacanthus horridus Bittner; p. 77, PI. 2 (fig. 1)
1895 - Periacanthus horridus Bittner in Bittner; p. 250, PI. 1
(figs. 1-2)
1898 - Periacanthus horridus Bittner in Lòrenthey; p. 34
1 899 - Periacanthus horridus Bittner in Oppenheim; p. 57
191 Oa — Periacanthus horridus Bittner in Fabiani; p. 22
1915 - Periacanthus horridus Bittner in Fabiani; p. 284
1929 - Periacanthus horridus Bittner in Glaessner; p. 31 1
1929 - Periacanthus horridus Bittner in Lòrenthey & Beurlen; p. 152,
PI. 7 (fig. 8)
1959 - Periacanthus horridus Bittner in Via Boada; p. 374
1969 - Periacanthus horridus Bittner in Via Boada; p. 175, PI. 12
(fig- 2)
1981 - Periacanthus horridus Bittner in Quayle & Collins; p. 744,
PI. 104 (fig. 14)
1989 - Periacanthus horridus Bittner in Solò & Via Boada;
p. 31
1982 - Periacanthus horridus Bittner in Busulini, Tessier & Visentin;
p. 78, Text-fig. 2
1994 - Periacanthus horridus Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 177, PI. 5 (fig. 1)
1995 - Periacanthus horridus Bittner in De Angeli; p. 14, Text-fig. 2
(4), PI. 1 (fig. 5)
2001 - Periacanthus horridus Bittner in De Angeli & Beschin; p. 26,
Text-fig. 20
2004 - Periacanthus horridus Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 115
2005 - Periacanthus horridus Bittner in Beschin, De Angeli, Checchi
& Zarantonello; p. 20
Holotype: unknown.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Bittner (1875, 1895) reported this species
from S. Giovanni Ilariore (Verona) (paratype: MNHB
K. 1 1 0, MB. A. 660); Busulini et al. ( 1 982) reported one
specimen from “Main” quarry of Arzignano (Vicenza)
(repository and catalogue number unknown); Beschin
et al. (1994) reported three specimens (MCZ 1264,
1416, 1448) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza); De Angeli (1995) reported one
specimen (MCZ 1495) from “Fontanella” di Grancona
(Vicenza).
Note: this species is also reported from thè middle/
upper Eocene of Spain, Great Britain, and Hungary.
Family Mithracidae Balss, 1929
Micippa Leach, 1817
Type species: Cancer cristatus Linnaeus, 1758.
Stratigraphic range: lower Oligocene (Rupelian)
Recent.
Micippa antiqua Beschin, De Angeli & Checchi, 2001
2001 - Micippa antiqua Beschin, De Angeli & Checchi; p. 1 8, Text-
fig. 5, PI. 2 (fig. 1 a-b)
2001 - Micippa antiqua Beschin, De Angeli & Checchi in De Angeli
& Beschin; p. 27
Holotype: MCZ 2122 (I.G. 286477).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported one specimen
(MCZ 2122) from Castelgomberto (Vicenza).
Mithracia Bell, 1858
Type species: Mithracia libinoides Bell, 1858.
Stratigraphic range: lower/middle Eocene (Ypresian-
Lutetian).
50
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Mithracia margaritifera Beschin, Busulini, De Angeli &
Tessier, 1994
1994 - Mithracia margaritifera Beschin, Busulini, De Angeli &
Tessier; p. 1 79, Text-fig. 5, PI. 6 (fig. 2 a-c)
2001 - Mithracia margaritifera Beschin, Busulini, De Angeli & Tessier
in De Angeli & Beschin; p. 27
Holotype: MCZ 1 296 (I.G. 211811).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported two
specimens (MCZ 1284, 1296) from “Boschetto” and
“Albanello” quarries of Nogarole Vicentino (Vi¬
cenza).
Mithracia oppionii Larghi, 2002
1998 - ? Mithracia sp. in Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 23, Text-figs. 9 (3), 1 1
2000 - ? Mithracia sp. in Beschin, De Angeli & Alberti; p. 1 5
2001 - 1 Mithracia sp. in De Angeli & Beschin; p. 27
2002 - Mithracia oppionii Larghi; p. 62, Text-figs. 2-3
Holotype: MCZ 2064 (I.G. 286342).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1998) reported three speci¬
mens (MCZ 1524, 1525, 1526) from “Rossi” quarry of
Monte di Malo (Vicenza); Larghi (2002) reported one
specimen (MCZ 2064) from “Lovara” quarry of Chiampo
(Vicenza).
Family Pisidae Dana, 1851
Pisa Leach, 1814
Type species: Cancer biaculeatus Montagu, 1813.
Stratigraphic range: Pleistocene - Recent.
Pisa armata (Latreille, 1803)
1803 - Maja armata Latreille; p. 98
1815 - Pisa gibbsi Leach; PI. 19 (Figs. 1-4), nov. syn.
1914 - Pisa gibbsi Leach in M. Gemmellaro; p. 84, PI. 1 (fig. 15)
1918 - Pisa armala (Latreille) in Pesta; p. 344, Text-figs. 110-111
1929 - Pisa armata (Latreille) in Glaessner; p. 318
1931 - Pisa armata (Latreille) in Nobre; p. 155, Text-figs. 91-94
1940 - Pisa gibbsi Leach in Bouvier; p. 331, Text-fig. 202, PI. 13
(fig- 5)
1 946 - Pisa gibbsi Leach in Zariquiey Alvarez; p. 170, PI. 23 (d)
1950 - Pisa gibbsi Leach in Zariquiey Alvarez; p. 109, PI. 5 (fig. 2)
1 956 - Pisa gibbsi Leach in Monod; p. 486, Text-fig. 654
1958 - Pisa armata (Latreille) in Holthuis & Gottlieb; p. 1 19
1961 - Pisa gibbsi Leach in Nunes-Ruivo; p. 31
1968 - Pisa armata (Latreille) in Zariquiey Alvarez; p. 454, Text-
figs. 151 d, 152 e, 154 d
1992 - Pisa armata (Latreille) in Falciai & Minervini; p. 251-252
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported one speci¬
men from Monte Pellegrino (Palermo) (MGUP, catalogue
number unknown).
Superfamily Parthenopoidea MacLeay, 1838
Family Dairidae Ng & Rodriguez, 1986
Daira De Haan, 1833
Type species: Cancer perlatus Herbst, 1790.
Stratigraphic range: middle Eocene (Lutetian) -
Recent.
Daira coronata Beschin, De Angeli, Checchi &
Zarantonello, 2005
2005 -Daira coronata Beschin, De Angeli, Checchi & Zarantonello;
p. 21, Text-fig. 13, PI. 4 (fig. 6)
Holotype: MCZ 2361 (I.G. 296580).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported one speci¬
men (MCZ 2361) from Grola di Comedo Vicentino
(Vicenza).
Daira depressa (A. Milne Edwards, 1865)
1865 - Phlyctenodes depressus A. Milne Edwards; p. 367, PI. 33
(figs. 2-2 b)
1 883 - Phlyctenodes depressus A. Milne Edwards in Bittner; p. 3 1 1
1896 - Pilumnus sp. in Ristori; p. 506, PI. 12 (fig. 10)
1905 - Phlyctenodes depressus A. Milne Edwards in Airaghi; p. 205,
PI. 4 (fig. 3)
191 Oa - Phlyctenodes depressus A. Milne Edwards in Fabiani;
p. 25
1915 - Phlyctenodes depressus A. Milne Edwards in Fabiani; p. 285
1929 - Phlyctenodes depressus A. Milne Edwards in Glaessner; 135
1969 - Daira depressa (A. Milne Edwards) in Via Boada; p. 373
2001 - Daira depressa (A. Milne Edwards) in Beschin, De Angeli &
Checchi; p. 20, PI. 2 (figs. 2-4)
2001 - Daira depressa (A. Milne Edwards) in De Angeli & Beschin;
p. 28
2004 - Daira depressa (A. Milne Edwards) in Beschin, Busulini, De
Angeli & Tessier; p. 1 15
2005 - Daira depressa (A. Milne Edwards) in Beschin, De Angeli,
Checchi & Zarantonello; p. 21, 22
Holotype: unknown.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: A. Milne Edwards (1862-1865) and Aira¬
ghi (1905) reported one specimen from Castelgom-
berto (Vicenza) (repository and catalogue number
unknown); Ristori (1896) reported one specimen
from S. Urbano (Vicenza) (repository and catalogue
number unknown); Beschin et al. (2001) reported 22
specimens (MCZ from 2095 to 2116), from Castel-
gomberto (Vicenza).
Daira eocenica var. sicula (Di Salvo, 1933)
1933 - Phymatocarcinus eocenicus var. sicula Di Salvo; p. 23, PI. 1
(fig. 5 a-d)
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DEC APODA FROM ITALY
51
2001 - Daira eocenica var. siculo (Di Salvo) in Beschin, De Angeli &
Checchi; p. 20
2005 - Daira eocenica var. siculo (Di Salvo) in Beschin, De Angeli,
Checchi & Zarantonello; p. 21, 22
Holotype: MGUP 2592.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported five speci-
mens (MGUP 2592, 2612, 2614, 2615, 2616) from
Balzo del Gatto, Bichinello, and La Pietra Lunga
(Palermo).
Daira salebrosa Beschin, Busulini, De Angeli & Tessier,
2002
2002 - Daira salebrosa Beschin, Busulini, De Angeli & Tessier; p. 1 5,
Text-fig. 10, PI. 2 (figs. 5-6)
2004 - Daira salebrosa Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 115
2005 - Daira salebrosa Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli, Checchi & Zarantonello; p. 21, 22, PI. 4
(fig- 5)
Holotype: MCZ 2271 (I.G. 296399).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported three speci-
mens (MCZ2271, 2272, 2273) from “Main” quarry of
Arzignano (Vicenza); Beschin et al. (2005) reported
three specimens (MCZ 2304, 2316, 2317) from Grola di
Comedo Vicentino (Vicenza).
Family Daldorfìidae Ng & Rodriguez, 1986
Daldorfia Rathbun, 1 904
Type species: Cancer horridus Linnaeus, 1754.
Stratigraphic range: Oligocene - Recent.
Daldorfia fabianii Beschin, De Angeli & Checchi,
2001
2001 - Daldorfia fabianii Beschin, De Angeli & Checchi; p. 22, Text-
fig. 6, PI. 2 (fig. 5)
2001 - Daldorfia fabianii Beschin, De Angeli & Checchi in De Angeli
& Beschin; p. 28
2004 - Daldorfia fabianii Beschin, De Angeli & Checchi in Beschin &
De Angeli; p. 21
Holotype: MCZ 2118 (I.G. 286473).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported one specimen
(MCZ 2118) from Castelgomberto (Vicenza).
Family Parthenopidae MacLeay, 1838
Parthenope Weber 1795
Parthenope angulifrons Latreille, 1825
1825 - Parthenope angulifrons Latreille; p. 46
1 863 - Lambrus angulifrons (Latreille) in Heller; p. 57, PI. 2
1918 - Lambrus angulifrons (Latreille) in Pesta; p. 371, Text-fig. 1 19
1931 - Lambrus angulifrons (Latreille) in Nobre; p. 140, Text-
figs. 80-81
1940 - Lambrus angulifrons (Latreille) in Bouvier; p. 310, Text-
fig. 191, PI. 12 (fig. 1)
1941 - Lambrus angulifrons (Latreille) in Zariquiey Cenarro; p. 351,
365, Text-figs. 36-44 a-c
1946 - Lambrus angulifrons (Latreille) in Zariquiey Alvarez; p. 167,
PI. 20
1968 - Parthenope angulifrons Latreille in Zariquiey Alvarez; p. 439,
Text-fig. 148 b
1992 - Parthenope angulifrons Latreille in Falciai & Minervini;
p. 242
2004b - Parthenope angulifrons Latreille in Garassino & De Angeli;
p. 22, Text-figs. 2, 4
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: Garassino & De Angeli (2004b) reported
one specimen (MSNM i 26283) from Favignana Island
(Sicilia).
Parthenope nummulitica (Bittner, 1875)
1875 - Lambrus nummuliticus Bittner; p. 79, PI. 1 (fig. 11)
191 Oa — Lambrus nummuliticus Bittner in Fabiani; p. 22
1915 - Lambrus nummuliticus Bittner in Fabiani; p. 284
1929 - Parthenope? nummulitica (Bittner) in Glaessner; p. 307
1983 - Parthenope nummulitica (Bittner) in Busulini, Tessier, Visentin,
Beschin, De Angeli & Rossi; p. 62, PI. 2 (fig. 2)
1994 - Parthenope nummulitica (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 182, PI. 6 (fig. 3)
2001 - Parthenope nummulitica (Bittner) in De Angeli & Beschin;
p. 28
2004 - Parthenope nummulitica (Bittner) in Beschin, Busulini,
De Angeli & Tessier; p. 1 1 5
2006 - Parthenope nummulitica (Bittner) in Beschin, De Angeli, Chec¬
chi & Mietto; p. 107, PI. 2 (fig. 8)
Holotype: MNHB K. 109, MB. A. 659.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Bittner (1875) reported one specimen
(MNHB K109, MB. A. 661) from S. Giovanni Barione
(Verona); Busulini et al. (1983) reported one specimen
(*28) from “Main” quarry of Arzignano (Vicenza);
Beschin et al. (1994) reported two specimens (MCZ
1307, 1433) from “Boschetto” quarry of Nogarole
Vicen-tino (Vicenza); Beschin et al. (2006) reported
one specimen (MCZ 2444) from Grotta della Poscola
(Vicenza).
Parthenope sp.
1 89 1 b -Lambrus sp. in Ristori; p. 21, PI. 1 (figs. 26-28)
Stratigraphic range: Pliocene.
Occurrence: Lazio.
Type species: Cancer longimanus Linnaeus, 1758.
Stratigraphic range: Eocene - Recent.
52
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: Ristori ( 1 89 1 b) reported some specimens
from Monte Mario, housed in Zuccari’s and Clerici’s col-
lections (repository and catalogne number unknown).
Parthenope sp.
1895 - Lambrus sp. in Crema; p. 674, Text-fig. 14
Stratigraphic range: Miocene.
Occurrence: Piemonte.
Material: Crema (1895) reported some specimens
from Torino, today lost.
Parthenope sp.
1914 - Lambrus sp. in M. Gemmellaro; p. 58, PI. 1 (figs. 16-17)
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported some spe¬
cimens from Monte Pellegrino and Ficarazzi (Palermo)
(MGUP, catalogne number unknown) and Marchese
di Monterosato’s collection (repository and catalogue
number unknown).
Parthenope sp.
2004a - Parthenope sp. in Garassino & De Angeli; p. 41, Text-figs. 3
(1-3), 4 (6)
Stratigraphic range: Pliocene, Pleistocene.
Occurrence: Emilia Romagna.
Material: Garassino & De Angeli (2004a) repor¬
ted four specimens (MG 0631, 0640, 0641, 0642)
from Fiume Arda (Piacenza) and Torrente Stirone
(Parma).
Superfamily Retroplumoidea Gill, 1894
Family Retroplumidae Gill, 1894
Loerenthopluma Beschin, Busulini, De Angeli &
Tessier, 1996
Type species: Loerenthopluma lata Beschin, Busulini,
De Angeli & Tessier, 1996.
Stratigraphic range: middle Eocene (Lutetian).
Loerenthopluma lata Beschin, Busulini, De Angeli &
Tessier, 1996
1996b - Loerenthopluma tata Beschin, Busulini, De Angeli & Tes¬
sier; p. 89, Text-fig. 3, PI. 1 (fig. 1)
1998 - Loerenthopluma lata Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli, Tessier & Ungaro; p. 31, Text-figs.
15(4), 17
2000 - Loerenthopluma lata Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli & Alberti; p. 15
2001 - Loerenthopluma lata Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 28, Text-fig. 21(1)
Holotype: MCZ 1476 (I.G. 286351).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1996b) reported two speci¬
mens (MCZ 1476, 1477) from “Rossi” quarry of Monte
di Malo (Vicenza).
Retrocypoda Via Boada, 1959
Type species: Retrocypoda almelai Via Boada, 1959.
Stratigraphic range: middle Eocene (Lutetian).
Retrocypoda almelai Via Boada, 1959
1959 - Retrocypoda almelai Via Boada; p. 394, Text-fig. 20
1969 - Retrocypoda almelai Via Boada in Via Boada; p. 331, Text-
fig. 41, PI. 38 (fig. 4); PI. 39 (figs. 1-5)
1980 - Retrocypoda almelai Via Boada in Via Boada; p. 8, PI. 1
(fig. 5)
1989 - Retrocypoda almelai Via Boada in Solè & Via Boada; p. 3 1
1996b - Retrocypoda almelai Via Boada in Beschin, Busulini,
De Angeli & Tessier; p. 96, Text-fig. 5, PI. 2 (fig. 1 a-c)
2001 - Retrocypoda almelai Via Boada in De Angeli & Beschin; p. 28,
Text-fig. 21 (3)
Holotype: MGSB 15.942.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1996b) reported one speci¬
men (MCZ 1475) from “Albanello” quarry of Nogarole
Vicentino (Vicenza).
Note: this species is also reported from thè Eocene of
Spain.
Retropluma Gill, 1 894
Type species: Archaeoplax notopus Alcock &Ander-
son, 1894.
Stratigraphic range: Eocene - Recent.
Retropluma craverii (Crema, 1895)
1895 - Goneplax craverii Crema; p. 675, Text-fig. 16
1969 - Retropluma craverii (Crema) in Via Boada; p.
1996b - Retropluma craverii (Crema) in Beschin, Busulini, De Angeli
& Tessier; p. 95
2004a - Retropluma craverii (Crema) in Garassino & De Angeli;
p. 45
2004 - Retropluma craverii (Crema) in Garassino, De Angeli, Gallo &
Pasini; p. 255
Holotype: MCC.
Stratigraphic range: upper Pliocene (Piacentian).
Occurrence: Piemonte.
Material: Crema (1895) reported one speci¬
men from Bra (Cuneo) (MCC, catalogue number
unknown).
Retropluma eocenica Via Boada, 1959
1959 - Retropluma eocenica V ia Boada; p. 392, Text-fig. 19
1969 - Retropluma eocenica Via Boada in Via Boada; p. 323, Text-
fig. 40, PI. 38 (figs. 1-3)
1980 - Retropluma eocenica Via Boada in Via Boada; p. 56, PI. 1
(fig. 3)
1 982 - Retropluma eocenica Via Boada in Via Boada; p. 1 8, Text-fig. 1
1 989 - Retropluma eocenica Via Boada in Solè & Via Boada; p. 3 1
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
53
1996b - Retropluma eocenica Via Boada in Beschin, Busulini,
De Angeli & Tessier; p. 92, Text-fig. 4, PI. 1 (figs. 2-4)
2001 - Retropluma eocenica Via Boada in De Angeli & Beschin; p. 29,
Text-figs. 21 (2), 22
2004 - Retropluma eocenica Via Boada in Beschin, Busulini,
De Angeli & Tessier; p. 1 1 5
Holotype: MGSB 20.101.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1996b) reported nine speci-
mens (MCZ from 1467 to 1474, 1478) from “Albanello”
quarry of Nogarole Vicentino (Vicenza) and “Main”
quarry of Arzignano (Vicenza).
Note: this species is also reported from thè Eocene of
Spain.
Retropluma cff. R. eocenica Via Boada, 1959
2003 - Retropluma cfr. R. eocenica Via Boada in Larghi; p. 59, Text-
figs. 2-3 a
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte.
Material: Larghi (2003) reported one specimen
(MSNM Ì26238) from Ponzone (Alessandria).
Superfamily Cancroidea Latreille, 1 802
Family Atelecyclidae Ortmann, 1893
Atelecyclus Leach, 1814
Type species: Cancer rotundatus Olivi, 1792.
Stratigraphic range: ?Eocene - Recent.
Atelecyclus elegans Ristori, 1896
1896 - Atelecyclus elegans Ristori; p. 508, PI. 12 (fig. 2)
1929 - Atelecyclus elegans Ristori in Glaessner; p. 62
1981 - Atelecyclus elegans Ristori in Delle Cave; p. 46
Holotype: IGF 839E.
Stratigraphic range: middle Miocene (Helvetian).
Locality: Piemonte.
Material: Ristori (1896) reported one specimen from
Albugnano (Torino).
Atelecyclus rotundatus (Olivi, 1792)
1792 - Cancer rotundatus Olivi; p. 47, PI. 2 (fig. 2)
1 813 - Cancer hippa septemdentatus Montagu; p. 1, PI. 1 (fig. 1)
1815 - Atelecyclus heterodon Leach; p. 2, Text-figs. 1-2
1863 - Atelecyclus heterodon Leach in Heller; p. 133
1888 - Atelecyclus heterodon Leach in Gourret; p. 55, PI. 1 (figs. 1-
17)
1905 - Atelecyclus rotundatus (Olivi) in Checchia-Rispoli; p. 86-98,
Text-fig. 1
1914 - Atelecyclus rotundatus var. checchiai M. Gemmellaro; p. 89,
PI. 1 (figs. 24-25), nov. syn.
1918 - Atelecyclus rotundatus (Olivi) in Pesta; p. 382, Text-fig. 122
1929 - Atelecyclus rotundatus var. checchiai M. Gemmallaro in
Glaessner; p. 62
1936 - Atelecyclus heterodon Leach in Nobre; p. 25, PI. 8 (fig. 14)
1928 - Atelecyclus septemdentatus Lebour; p. 524, Text-figs. 1 (5),
4 (24, 26-27), 5 (13-14), PI. 2 (fig. 2), PI. 9 (figs. 1-6), PI. 10
(figs. 1-2)
1940 - Atelecyclus septemdentatus Lebour in Bouvier; p. 219, Text-
fig. 148, PI. 8 (fig. 6)
1946 - Atelecyclus septemdentatus Lebour in Zariquiey Alvarez;
p. 149, PI. 10 (d-e)
1 956 - Atelecyclus septemdentatus Lebour in Monod; p. 148
1957 - Atelecyclus rotundatus (Olivi) in Forest; p. 472
1967 - Atelecyclus rotundatus (Olivi) in Alien; p. 29, 68, 102
1968 - Atelecyclus rotundatus (Olivi) in Zariquiey Alvarez; p. 342,
Text-fig. 1 1 2 b
1969 - Atelecyclus rotundatus (Olivi) in Christiansen; p. 37, Text-
fig. 13 a
1976 - Atelecyclus rotundatus (Olivi) in Turkey; p. 37
1980 - Atelecyclus rotundatus (Olivi) in Ingle; p. 103, Text-fig. 46,
PI. 14 a
1992 - Atelecyclus rotundatus (Olivi) in Falciai & Minervini; p. 196
2004b - Atelecyclus rotundatus (Olivi) in Garassino & De Angeli;
p. 23, Text-figs. 3, 5
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1905) reported one spe¬
cimen from Monte Pellegrino (Palermo) (MGUP, cata¬
logne number unknown); Garassino & De Angeli (2004b)
reported one specimen (MSNM Ì26282) from Favignana
Island (Sicilia).
Family Cancridae Latreille, 1802
Cancer Linnaeus, 1758
Type species: Cancer pagurus Latreille, 1810.
Stratigraphic range: Miocene - Recent.
Cancer spinosus (Ristori, 1886)
1886 - Pilumnus spinosus Ristori; p. 104, PI. 2 (fig. 8)
1 89 1 b - Pilumnus spinosus Ristori in Ristori; p. 20
1892b - Pilumnus spinosus Ristori in Ristori; p. 90
1929 - Cancer spinosus (Ristori) in Glaessner; p. 108
Holotype: lost.
Stratigraphic range: upper Pliocene (Piacentian).
Occurrence: Lazio, Sicilia.
Material: Ristori (1886) reported one specimen from
Tremonte (Sicilia) today lost; Ristori ( 1 89 1 b) reported
one specimen from Monte Mario (Roma), housed in Zuc-
cari’s collection (MGPUR, catalogue number unknown);
Ristori (1892b) reported some specimens from thè Piano¬
sa Island (repository and catalogue number unknown).
Ceronnectes De Angeli & Beschin, 1998
Type species: Cancer bòckhi Lòrenthey, 1897.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Ceronnectes boeckhi (Lòrenthey, 1 897)
1897 - Cancer Bòckhi Lòrenthey; p. 99
54
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1 898 - Cancer Bóckhi Lòrenthey in Lòrenthey; p. 57, PI. 4 (fig. 5)
1929 - Necronectes Bóckhi (Lòrenthey) in Lòrenthey & Beurlen;
p. 168, PI. 8 (fig. 8)
1 929 - Necronectes Bóckhi (Lòrenthey) in Glaessner; p. 265
1998 - Ceronnectes boeckhi (Lòrenthey) in De Angeli & Beschin;
p. 89, Text-figs. 1-2
2001 - Ceronnectes boeckhi (Lòrenthey) in De Angeli & Beschin;
p. 32
2004 - Ceronnectes boeckhi (Lòrenthey) in Beschin, Busulini,
De Angeli & Tessier; p. 1 15
Holotype: unknown
Stratigraphic range: middle/upper Eocene.
Occurrence: Veneto.
Material: De Angeli & Beschin (1998) reported one
specimen (MCZ 1563) from “Main” quarry of Arzignano
(Vicenza).
Note: Ceronnectes boeckhi must be included in thè
family Cancridae Latreille, 1802, as pointed out by
Lòrenthey (1897). This species is also reported from thè
upper Eocene of Hungary. Anisospinos Schweitzer &
Feldmann, 2000, has a carapace with five evenly-spaced
frontal spine including inner-orbital spine, orbits broad,
deeply excavated, with well-developed rim; anterolateral
margin with eight spines including outer-orbital spine,
spines separated to bases; third and fourth, fifth and sixth,
and seventh and eighth anterolateral spines paired; pos-
terolateral margin entire, rim weak if present; carapace
regions distinct, granular or omamented with tubercles.
Anisospinos and Ceronnectes have shape and morpho-
logical characters of thè carapace very similar. The only
difference between thè two genera is thè more (in Anisos¬
pinos) or less (in Ceronnectes ) development of thè frontal
and inner-orbital spine of Anisospinos. This difference
must be considered as infraspecific and therefore Anisos¬
pinos could be considered as synonym with Ceronnectes.
Lobocarcinus Reuss, 1857
Type species: Cancer paulinowurttembergensis v.
Meyer, 1847.
Stratigraphic range: Miocene - Recent.
Lobocarcinus sismondai (v. Meyer, 1843)
1 822 - Cancer punctulatus Desmarest; p. 92, PI. 7 (figs. 3-4)
1839 - Cancer punctulatus Desmarest in A. Sismonda; p. 93, Text-
figs. A-B
1843 - Cancer sismondae v. Meyer; p. 590
1846 - Platycarcinus antiquus E. Sismonda; p. 58, PI. 3 (figs. 1-2),
nov. syn.
1857 - Platycarcinus antiquus E. Sismonda in Meneghini; p. 528,
PI. H (fig. 11)
1858 - Lobocarcinus sismondai (v. Meyer) in Reuss; p. 41, PI. 9
(figs. 1-2)
1858 - Lobocarcinus imperator Reuss; p. 41, Pls. 7-8, PI. 9 (fig. 1)
1861 - Platycarcinus sismondae (v. Meyer) in E. Sismonda; p. 18
1861 - Platycarcinus deshayesii A. Milne Edwards; p. 88
1861 - Cancer sismondai v. Meyer in Michelotti; p. 140
1864 - Cancer sismondae (v. Meyer) in A. Milne Edwards; p. 316,
Pls. 24-25
1864 - Cancer deshayesii (A. Milne Edwards) in A. Milne Edwards;
p. 314, PI. 22 (figs. 1-2), PI. 23 (fig. 1)
1875 - Platycarcinus sismondai (v. Meyer) in Bittner; p. 23
1886 - Cancer sismondae (v. Meyer) in Ristori; p. 95, PI. 2 (fig. 1)
1887 - Cancer sismondae (v. Meyer) in Mariani & Parona; p. 152
1889 - Cancer sismondae (v. Meyer) in Ristori; p. 217
1 89 1 a - Cancer sismondae (v. Meyer) in Ristori; p. 4
1893 - Cancer cfr. illyricus in Bittner; p. 32
1895 - Cancer sismondae (v. Meyer) in Crema; p. 679, Text-
fig. 19 a-c
1 896 - Platycarcinus sismondai (v. Meyer) in Vinassa de Regny;
p. 124, PI. 2 (fig. 1 a-b)
1 903 - Cancer sismondae (v. Meyer) in Lòrenthey; p. 32
1907 - Cancer sismondae (v. Meyer) in Lòrenthey; p. 204, 208, 210
1908? - Cancer sismondae (v. Meyer) in Couffon; p. 5, PI. 2
(figs. 3-4)
1908? - Cancer deshayesii (A. Milne Edwards) in Couffon; p. 5,
PI. 2 (fig. 11)
1909 - Cancer sismondae (A. Milne Edwards) in Lòrenthey; p. 240
191 Oa — Cancer sismondae (v. Meyer) in Fabiani; p. 33
1921 - Cancer deshayesii (A. Milne Edwards) in Bell; p. 7
1924 - Cancer ( Lobocarcinus ) sismondae (v. Meyer) in Glaessner;
p. 115
1927 - Cancer deshayesii (A. Milne Edwards) in Van Straelen; p. 87,
PI. 3 (fig. 2), PI. 4 (figs. 1-2)
1929 - Cancer sismondai (v. Meyer) in Glaessner; p. 106
1929 - Cancer sismondai (v. Meyer) in Lòrenthey; p. 161
1934 - Cancer deshayesii (A. Milne Edwards) in Van Straelen;
p. 207
1934 - Cancer sismondai (v. Meyer) in Van Straelen; p. 207
1946 - Cancer sismondai var. antiatina, Maxia; p. 134, Text-fig. 1,
PI. 1 (figs. 2-5), nov. syn.
1950 - Cancer sismondae (v. Meyer) in Comaschi Caria; p. 326
1950 - Platycarcinus antiquus E. Sismonda in Comaschi Caria;
p. 327
1956 - Cancer sismondai (v. Meyer) in Comaschi Caria; p. 282, 284, 288
1956 - Platycarcinus antiquus E. Sismonda in Comaschi Caria;
p. 283
1961 - Cancer sismondai var. antiatina Maxia in Zappi; p. 86
1965 - Cancer sismondai (v. Meyer) in Varola; p. 295
1965 - Cancer sismondai var. antiatina Maxia in Giannelli, Salvatorini
& Tavani; p. 521
1969 - Cancer sismondai var. antiatina Maxia in Largaiolli, Martinis,
Nardin, Rossi & Ungaro; p. 29
1969 - Cancer sismondai var. antiatina Maxia in Rossi; p. 22
1977 - Cancer cfr. sismondai (v. Meyer) in Georgiades Dikeoulia;
p. 420
1981 - Cancer sismondai (v. Meyer) in Varola; p. 16, PI. 3 (figs. 5-6),
PI. 4 (figs. 1-2), PI. 5 (figs. 1-2), PI. 6 (figs. 2, 4)
1982 - Cancer sismondai (v. Meyer) in Via Boada, Martinell &
Domènech; p. 242, Pls. 1-2
1982 - Cancer sismondai (v. Meyer) in Bonfiglio & Donadeo; p. 270,
Text-figs. 5-27, Pls. 33-44
1982 - Cancer sismondai (v. Meyer) in Bonfiglio; p. 5, Text-figs. 1-4,
Pls. 1-7
1984 - Cancer sismondai (v. Meyer) in Miiller; p. 75
1988 - Cancer sismondai (v. Meyer) in Solé & Via Boada; p. 35
1990 - Cancer sismondai (v. Meyer) in Moisette & Miiller; p. 739,
PI. 1 (fig. 1), PI. 2 (figs. 1-2)
1993 - Cancer sismondai (v. Meyer) in Miiller; p. 14
1997 - Cancer sismondai (v. Meyer) in Beschin & Santi; p. 13, PI. 1
(figs. 1-2)
1998 - Cancer sismondai (v. Meyer) in Miiller; p. 28
2000 - Cancer sismondai (v. Meyer) in Garassino & Fomaciari; p. 29,
Text-fig. 1
2000 - Lobocarcinus sismondai (v. Meyer) in Schweitzer & Feldmann;
p. 244
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
55
2002 - Lobocarcinus sismondai (v. Meyer) in Collins; p. 86
2002 - Lobocarcinus sismondae (v. Meyer) in Bortoluzzi; p. 17, Text-
fig. 7
2004a - Lobocarcinus sismondae (v. Meyer) in Garassino & De Angeli;
p. 42, Text-figs. 9-11
;J1
Holotype: unknown.
Stratigraphic range: Miocene - Pleistocene.
Occurrence: Piemonte, Emilia Romagna, Lazio,
Puglia, Calabria, Sardegna, Sicilia.
Material: Meneghini (1857) reported some speci-
mens from Capo S. Marco (Oristano) (repository and cat-
alogue number unknown); A. Sismonda (1839) reported
two specimens from S. Stefano Roero (Asti), today lost;
E. Sismonda (1846) reported one specimen from Qua-
glina (Asti), housed in thè Sotteri’s collection, today
lost; A. Milne Edwards (1861) reported one specimen
(MNHN A24543) from Alba; Ristori (1886) reported
some specimens from Quaglina, S. Stefano Roero and
Verruca Savoja (Asti), Monte Capriolo (Bra) (repository
and catalogue number unknown), four specimens from
Fornaci (Savona), today lost, and some specimens from
Tremonte (Sicilia) (repository and catalogue number
unknown); Ristori (189 la) reported one specimen from
Anzio Tor Caldara (Roma), housed in Clerici’s collec¬
tion (MGPUR, catalogue number unknown); Crema
(1895) reported some specimens from Bra (repository
and catalogue number unknown); Vinassa de Regny
( 1 896) reported one specimen from Lesignano dei Bagni
(Parma) (MGPUP, catalogue number unknown); Lòren-
they (1909) reported some specimens from Cagliari
(Sardegna), housed in Lovisato’s collection (repository
and catalogue number unknown); Maxia (1946) reported
one specimen from Anzio (Roma), housed in Clerici’s
collection today lost; Comaschi Caria (1950) reported
this species from Capo S. Marco (Oristano) (repository
and catalogue number unknown); Varola (1965) reported
13 specimens from Porto Graulo (Lecce) (repository and
catalogue number unknown); Varola (1981) reported
102 specimens from Porto Craulo (Otranto) (MPUP and
GNSL, catalogue number unknown); Bonfiglio & Dona-
deo (1982) reported 28 specimens from Torre dell’Orso
(Lecce) (MPM, catalogue number unknown); Beschin
& Santi (1997) reported four specimens (MCV A, B;
MCZ 1572, 1573) from Vignola (Modena); Garassino
& Fomaciari (2000) reported ten specimens from Fiume
Enza (Parma), housed in a private collection (catalogue
number unknown); Bortoluzzi (2002) reported one
specimen (MGPD 28976) from Strongoli (Catanzaro);
Garassino & De Angeli (2004a) reported eight speci¬
mens (MG 0564, 0565, 0613, 0619, from 0615 to 0618)
from Fiume Arda (Piacenza), Fiume Enza, and Campore
quarry (Parma).
Note: this species is also reported from thè Miocene-
Pleistocene of Great Britain, thè Netherlands, Spain, Hun-
gary, Greece, Austria, and Algeria.
Family Cheiragonidae Ortmann, 1893
Montezumella Rathbun, 1930
Type species: Montezumella tabulata Rathbun, 1930.
Stratigraphic range: Eocene - Miocene.
Montezumella elegans (Lòrenthey & Beurlen, 1 929)
1929 - Titanocarcinus elegans Lòrenthey & Beurlen; p. 235, PI. 1 1
(fig- 3)
1981 - Montezumella elegans (Lòrenthey & Beurlen) in Quayle &
Collins; p. 748
1995 - Montezumella elegans (Lòrenthey & Beurlen) in De Angeli;
p. 14, Text-fig. 3 (1), PI. 1 (fig. 7), PI. 2 (fig. 1)
2001 - Montezumella elegans (Lòrenthey & Beurlen) in De Angeli &
Beschin; p. 29, Text-fig. 23
Holotype: unknown.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli (1995) reported two specimens
(MCZ 1496, 1497), from “Fontanella” di Grancona
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Montezumella pumicosa Beschin, Busulini, De Angeli &
Tessier, 2002
2002 - Montezumella pumicosa Beschin, Busulini, De Angeli &
Tessier; p. 16, Text-fig. 11, PI. 3 (fig. 1 a-b)
2004 - Montezumella pumicosa Beschin, Busulini, De Angeli & Tessier
in Beschin, Busulini, De Angeli & Tessier; p. 1 1 5
Holotype: MCZ 2274 (I.G. 296402).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported one speci¬
men (MCZ 2274) from “Main” quarry of Arzignano
(Vicenza).
Montezumella scabra Quayle & Collins, 1981
1981 - Montezumella scabra Quayle & Collins; p. 747, PI. 105 (fig. 1)
1983 - Montezumella cfr. scabra Quayle & Collins in Busulini, Tessier,
Visentin, Beschin, De Angeli & Rossi; p. 63, PI. 3 (fig. 6)
1994 - Montezumella scabra Quayle & Collins in Beschin, Busulini,
De Angeli & Tessier; p. 183, PI. 6 (fig. 4)
2001 - Montezumella scabra Quayle & Collins in De Angeli &
Beschin; p. 29
Holotype: NHM 61711.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Busulini et al. (1983) reported four speci¬
mens (*143, *8, *144; MSNV 10093), from “Main”
quarry of Arzignano (Vicenza); Beschin et al. (1994)
reported one specimen (MCZ 1287), from “Boschetto”
quarry of Nogarole Vicentino (Vicenza).
Note: this species is also reported in thè upper Eocene
of Great Britain.
Superfamily Portunoidea Rafìnesque-Schmaltz, 1815
Family Geryonidae Colosi, 1924
Coeloma A. Milne Edwards, 1865
Type species: Coeloma vigil A. Milne Edwards, 1865.
56
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: Eocene - Oligocene.
Coe/oma isseli Ristori, 1886
1886 - Coeloma isseli Ristori; p. 119, PI. 2 (fig. 10)
1923 - Coeloma isseli Ristori in Tettoni; p. 161
1929 - Coeloma sp. in Glaessner; p. 122
1929 - Coeloma ? isseli Ristori in Glaessner; p. 120
1981 - Coeloma isseli Ristori in Delle Cave; p. 48
Syntype: IGF 959E (fìgured by Ristori in PI. 2, fig. 10).
Stratigraphic range: Pliocene.
Occurrence: Emilia Romagna.
Material: Ristori (1886) reported some specimens
(IGF from 960E to 97 1E, syntypes) from S. Venanzio
(Modena); Tettoni (1923) reported one specimen from
S. Venanzio (Modena) (MGUM, catalogue number unk-
nown).
Coeloma sabatium Ristori, 1886
1886 - Coeloma sabatium Ristori; p. 117, PI. 2 (figs. 11-12, 17)
1887 - Coeloma rupeliense Stainier; p. 87
1929 - Coeloma sabatium Ristori in Glaessner; p. 121
Holotype: lost.
Stratigraphic range: Pliocene.
Occurrence: Liguria.
Material: Ristori (1886) reported one specimen from
Fornaci (Savona), housed in don Perrando’s collection,
today lost.
Coeloma vigil A. Milne Edwards, 1865
1861 - Cancer beggiatoi Michelotti; p. 140, PI. 14 (figs. 1-2)
1865 - Coeloma vigil A. Milne Edwards; p. 352, PI. 35 (figs. 1-3)
1875 - Coeloma vigil A. Milne Edwards in Bittner; p. 97, PI. 5
(figs. 1-4)
1883 - Coeloma vigil A. Milne Edwards in Bittner; p. 314
1889 - Coeloma vigil A. Milne Edwards in Ristori; p. 403, PI. 15
(figs. 4-5)
1901 - Coeloma vigil A. Milne Edwards in Oppenheim; p. 283
191 Oa - Coeloma vigil A. Milne Edwards in Fabiani; p. 33
1915 - Coeloma vigil A. Milne Edwards in Fabiani; p. 285
1929 - Coeloma vigil A. Milne Edwards in Glaessner; p. 122
1929 - Coeloma vigil A. Milne Edwards in Lòrenthey & Beurlen;
p. 243, PI. 15 (fig. 16)
1969 - Coeloma ( Coeloma ) vigil A. Milne Edwards in Glaessner; 524,
Text-fig. 332 (2 a)
1974 - Coeloma vigil A. Milne Edwards in Mastrorilli; p. 4, Text-
fig. without number
1987 - Coeloma ( Coeloma ) vigil A. Milne Edwards in Allasinaz;
p. 542, Text-fig. 12, PI. 6 (figs. 1-4), PI. 7 (figs. 1-5)
2001 - Coeloma ( Coeloma ) vigil A. Milne Edwards in De Angeli &
Beschin; p. 29, Text-fig. 24
2004 - Coeloma ( Coeloma ) vigil A. Milne Edwards in Garassino,
De Angeli, Gallo & Pasini; p. 270
Holotype: MNHN R03822.
Stratigraphic range: upper Eocene (Priabonian) -
lower Oligocene (Rupelian).
Occurrence: Piemonte, Liguria, Veneto.
Material: A. Milne Edwards (1862-1865) reported
two specimens (MNHN R03822, R03794) from thè
“nummulitico” of Vicenza; Bittner (1875) reported this
species from Laverda (Vicenza) (repository and catalogue
number unknown); Ristori ( 1 889) reported two specimens
from Sassello (Savona), housed in don Perrado’s collec¬
tion today in “DIP.TE.RIS” (we identified one originai
specimen: 2389/Sa-II-S 1 86 (ex Sa-II-S205), fìgured in
PI. 15, figs. 4-5); Allasinaz (1987) reported 30 specimens
from Ciglione, Ponzone, Fontanino di Rio Caramagna,
Rio Volpina and Case Cherpione (MCSN, catalogue
number unknown); Garassino et al. (2004) reported three
specimens (MSNM from Ì26279 to Ì26281) from Case
Cherpione (Ovada).
Note: this species is also reported from thè Oligocene
of Hungary.
Family Portunidae Rafinesque-Schmaltz, 1815
Boschettia Busulini, Tessier, Beschin & De Angeli, 2003
Type species: Boschettia giampietroi Busulini, Tessi¬
er, Beschin & De Angeli, 2003.
Stratigraphic range: middle Eocene (Lutetian).
Boschettia giampietroi Busulini, Tessier, Beschin &
De Angeli, 2003
1994 - Coeloma (Paracoeloma) sp. in Beschin, Busulini, De Angeli &
Tessier; p. 190, Text-fig. 7, PI. 9 (fig. 6)
2003 - Boschettia giampietroi Busulini, Tessier, Beschin &
De Angeli; p. 15, Text-figs. 2-4
Holotype: MCZ 2401 (I.G. 296785).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported one speci¬
men (MCZ 1447) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza); Busulini et al. (2003) reported two
specimens (MCZ 1447, 2401) from “Boschetto” quarry of
Nogarole Vicentino (Vicenza).
Carcinus Leach, 1814
Type species: Cancer maenas Linnaeus, 1758.
Stratigraphic range: Eocene - Recent.
Carcinus sp.
2004a - Carcinus sp. in Garassino & De Angeli; p. 44, Text-fig. 8
Stratigraphic range: Pliocene.
Occurrence: Emilia Romagna.
Material: Garassino & De Angeli (2004a) reported
four specimens (MG from 0643 to 0646) from Campore
quarry (Parma).
Carcinus sp.
2004 - Carcinus sp. in Garassino, De Angeli, Gallo & Pasini; p. 269
Stratigraphic range: Pliocene.
Occurrence: Piemonte.
Material: Garassino et al. (2004) reported four speci¬
mens (PU from 41 176 to 41179) from Cossato (Biella).
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
57
Charybdis De Haan, 1833
Type species: Cancer feriatus Linnaeus. 1758.
Stratigraphic range: Oligocene - Recent.
Charybdis antiqua (A. Milne Edwards, 1860)
1860 - Goniosoma antiqua A. Milne Edwards; p. 265, PI. 5 (fig. 4)
1875 - Goniosoma antiqua A. Milne Edwards in Bittner; p. 83
191 Oa — Charybdis antiqua (A. Milne Edwards) in Fabiani; p. 24
1915 - Cha/ybdis antiqua (A. Milne Edwards) in Fabiani; p. 285
1929 - Charybdis antiqua (A. Milne Edwards) in Glaessner; p. 1 13
2001 - Charybdis antiqua (A. Milne Edwards) in De Angeli &
Beschin; p. 31
Holotype: unknown.
Stratigraphic range; lower Oligocene (Rupelian).
Occurrence: Veneto.
Material; A. Milne Edwards (1862-1865) reported
this species from Salcedo (Vicenza) (repository and cata¬
logne number unknown).
Enoplonotus A. Milne Edwards, 1 860
Type species: Enoplonotus armatus A. Milne
Edwards, 1860.
Stratigraphic range: Eocene.
Enoplonotus armatus A. Milne Edwards, 1 860
1860 - Enoplonotus armatus A. Milne Edwards; p. 247, PI. 7
(figs. 1, 1 a)
1915 - Enoplonotus armatus A. Milne Edwards in Fabiani; p. 284
1929 - Enoplonotus armatus A. Milne Edwards in Glaessner; p. 149
1975 - Enoplonotus armatus A. Milne Edwards in Secretan; p. 359,
PI. 21 (figs. 1-2)
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: A. Milne Edwards ( 1 862- 1 865) reported one
specimen (? Massalongo’s collection) from Monte Bolca
(Verona) (repository and catalogue number unknonw);
Secretan (1975) reported two specimens (MSNV MI,
M2) from Monte Bolca (Verona).
Eocharybdis Beschin, Busulini, De Angeli & Tessier,
2002
Type species: Eocharybdis cristata Beschin, Busulini,
De Angeli & Tessier, 2002.
Stratigraphic range: middle Eocene (Lutetian).
Eocharybdis cristata Beschin, Busulini, De Angeli &
Tessier, 2002
2002 - Eocharybdis cristata Beschin, Busulini, De Angeli & Tessier;
p. 17, Text-fig. 12, PI. 3 (fig. 2)
2004 - Eocharybdis cristata Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 115
Holotype: MCZ 2275 (I.G. 296403).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported one speci¬
men (MCZ 2275) from “Main” quarry of Arzignano
(Vicenza).
Liocarcinus Stimpson, 1 87 1
Type species: Portunus holsatus Fabricius, 1798.
Stratigraphic range: Oligocene - Recent.
Liocarcinus cfr. L. rakosensis (Lòrenthey & Beurlen,
1929)
2004 - Liocarcinus cfr. L. rakosensis (Lòrenthey & Beurlen) in
Garassino, De Angeli, Gallo & Pasini; p. 269, Text-fig. 1 1
Stratigraphic range: upper Miocene (Messinian).
Occurrence: Piemonte.
Material: Garassino et al. (2004) reported one speci¬
men (PU 41198) from Cocconato (Asti).
Note: this species is also reported from thè Miocene
of Hungary.
Macropipus Prestandrea, 1833
Type species: Portunus macropipus Prestandrea, 1 833.
Stratigraphic range: Eocene - Recent.
Macropipus ovalipes Secretan, 1975
1975 - Macropipus ovalipes Secretan; p. 348, Text-figs. 12-18,
Pls. 17-20
Holotype: MSNV es. 8.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported one specimen
(MSNV es. 8) from Monte Bolca (Verona).
Necronectes A. Milne Edwards, 1881
Type species: Necronectes vidalianus A. Milne
Edwards, 1881.
Stratigraphic range: Oligocene - Miocene.
Necronectes schaff eri Glaessner, 1928
1928 - Necronectes schafferi Glaessner; p. 179, Text-fig. 3, PI. 3
(fig. 6)
1929 - Scylla cf. michelini A. Milne Edwards in Glaessner; p. 1 84
1929 - Scylla sp. cf. michelini A. Milne Edwards in Lòrenthey & Beur¬
len; p. 178, PI. 15 (figs. 5-6)
1929 - Necronectes schafferi Glaessner in Glaessner; p. 265
1933 - Necronectes schafferi Glaessner in Glaessner; p. 3, PI. 1
(fig- 1)
1992 - Necronectes schafferi Glaessner in De Angeli & Marangon;
p. 177, PI. 1 (figs. 1-2), PI. 2 (fig. 1 a-b)
1993 - Necronectes batalleri Via Boada in Miiller; p. 30, PI. 2
(fig- 3)
1998 - Necronectes schafferi Glaessner in Miiller; p. 30
Holotype: MHMW 1927/1/1.
Stratigraphic range: lower/middle Miocene (Burdi-
galian - Langhian).
Occurrence: Sardegna.
58
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: De Angeli & Marangon (1992) reported one
specimen (MCZ 1362) from Tresnuraghes (Oristano).
Note: this species is also reported from thè Miocene of
Austria, Hungary, Poland, Spain, and Malta.
Neptocarcinus Lòrenthey, 1897
Type species: Neptocarcinus millenaris Lòrenthey,
1897.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Neptocarcinus millenaris Lòrenthey, 1 897
1897 - Neptocarcinus millenaris Lòrenthey; p. 156
1898 - Neptocarcinus millenaris Lòrenthey in Lòrenthey; p. 179,
PI. 4 (fig. 3)
1929 - Neptocarcinus millenaris Lòrenthey in Lòrenthey & Beurlen;
p. 216, PI. 10 (fig. 3)
1983 - Neptocarcinus millenaris Lòrenthey in Busulini, Tessier, Visen¬
tin, Beschin, De Angeli & Rossi; p. 66, PI. 3 (fig. 3)
1991 - Neptocarcinus millenaris Lòrenthey in Miiller & Collins; p. 70,
Text-fig. 4 a, PI. 4 (fig. 11)
2001 - Neptocarcinus millenaris Lòrenthey in De Angeli & Beschin;
p. 32
2004 - Neptocarcinus millenaris Lòrenthey in Beschin, Busulini,
De Angeli & Tessier; p. 1 15
2005 - Neptocarcinus millenaris Lòrenthey in Beschin, De Angeli,
Checchi & Zarantonello; p. 22, Text-fig. 14, PI. 4 (fig. 7)
Holotype: MAFI.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Busulini et al. (1983) reported five speci-
mens (*1, *2, *3, *23; MSNVE 10096) from “Main”
quarry of Arzignano (Vicenza); Beschin et al. (2005)
reported ten specimens (MCZ 2300, 2301, 2351, 2352,
2374, 2375, 2376, 2380; MV 54, 04/15) from Grola di
Comedo Vicentino (Vicenza).
Note: this species is also reported from thè upper
Eocene of Hungary.
Nogarolia Beschin, Busulini, De Angeli & Tessier,
1994
Type species: Nogarolia mirabilis Beschin, Busulini,
De Angeli & Tessier, 1994.
Stratigraphic range: middle Eocene (Lutetian).
Nogarolia mirabilis Beschin, Busulini, De Angeli &
Tessier, 1994
1994 - Nogarolia mirabilis Beschin, Busulini, De Angeli & Tessier;
p. 184, Text-fig. 6, PI. 7 (fig. 1 a-b)
2001 - Nogarolia mirabilis Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 32, Text-fig. 26
Holoty pe: MCZ 1420 (I.G. 284609).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported one speci¬
men (MCZ 1420) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza).
Portunites Bell, 1858
Type species: Portunites incertus Bell, 1858.
Stratigraphic range: Eocene - Miocene.
Portunites rosenfeldi De Angeli & Garassino, 2006
2006a - Portunites rosenfeldi De Angeli & Garassino; p. 285, Text-
figs. 9 (a-b), 10
Holotype: MFSN 28995.
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Friuli-Venezia Giulia.
Material: De Angeli & Garassino (2006a) reported
two specimens (MFSN 28995, 29046) from Almadis
(Pordenone).
Portun us Weber, 1795
Type species: Cancer pelagicus Linnaeus, 1758.
Stratigraphic range: Eocene - Recent.
Portunus arcuatus (A. Milne Edwards, 1 860)
1860 - Neptunus arcuatus A. Milne Edwards; p. 240, PI. 6 (figs. 2,
2 a-b)
1875 - Neptunus arcuatus A. Milne Edwards in Bittner; p. 80
191 Oa — Neptunus arcuatus A. Milne Edwards in Fabiani; p. 23
1915 - Neptunus arcuatus A. Milne Edwards in Fabiani; p. 285
1929 - Neptunus arcuatus A. Milne Edwards in Glaessner; p. 267
1996 - Portunus arcuatus (A. Milne Edwards) in Beschin, Checchi &
Ungaro; p. 16
2001 - Portunus ( Portunus ) arcuatus (A. Milne Edwards) in De Angeli
& Beschin; p. 30
Holotype: unknown
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: A. Milne Edwards (1860) reported this spe¬
cies from Salcedo (Vicenza) (repository and catalogue
number unknown).
Portunus convexus (Ristori, 1889)
1889 - Neptunus convexus Ristori; p. 400, PI. 15 (fig. 1)
1929 - Neptunus convexus Ristori in Glaessner; p. 267
1950 - Neptunus convexus Ristori in Comaschi Caria; p. 326
1956 - Neptunus convexus Ristori in Comaschi Caria; p. 288
1965 - Neptunus convexus Ristori in Varola; p. 296
1969 - Neptunus convexus Ristori in Via Boada; p. 215
1974 - Neptunus convexus Ristori in Mastrorilli; p. 4
1987 - Portunus convexus (Ristori) in Allasinaz; p. 535, Text-fig. 10,
PI. 4 (figs. 3-8)
2003a - Portunus convexus (Ristori) in De Angeli & Marangon; p. 101,
Text-fig. 1 (3)
Holotype: lost.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Liguria, Piemonte, Sardegna.
Material: Ristori (1889) reported one specimen from
Sassello (Savona), housed in don Perrando’s collection, today
lost; Allasinaz (1987) reported 15 specimens from Ciglione,
Rio Caramagna, Rio Volpina, Case Cherpione and Ponzone
(Acqui) (PU; MCSN, catalogue number unknown).
CATALOGANO BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
59
Portunus cfr. P convexus (Ristori, 1889)
1 892c - Neptunus cfr. convexus (Ristori) in Ristori; p. 162
1 929 - Neptunus cfr. convexus (Ristori) in Glaessner; p. 267
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: Ristori (1892c) reported one specimen from
Chiavon (repository and catalogue number unknown).
Portunus edwardsi E. Sismonda, 1 846
1846 -Portunus edwardsi E. Sismonda; p. 70, PI. 3 (fig. 9)
1861 - Portunus edwardsi E. Sismonda in E. Sismonda; p. 20
1 886 - Portunus edwardsi E. Sismonda in Ristori; p. 1 10
189 la -Portunus edwardsi E. Sismonda in Ristori; p. 9, PI. 1 (fig. 13)
1929 -Portunus edwardsi E. Sismonda in Glaessner; p. 334
Holotype: lost.
Stratigraphic range: Pliocene.
Occurrence: Piemonte, Toscana.
Material: E. Sismonda (1846) reported one specimen
from Asti, today lost; E. Sismonda (1861) reported some
specimens from Asti, today lost; Ristori (1886) reported
some specimens from Asti, today lost; Ristori (189 la)
reported one specimen from Spicchio (Empoli), today
lost.
Portunus hastatus (Linnaeus, 1767)
1767 - Cancer hastatus Linnaeus; p. 1046
1863 - Lupa bastata (Linnaeus) in Heller; p. 77, PI. 2 (fig. 10)
1918 -Neptunus hastatus (Linnaeus) in Pesta; p. 411, Text-fig. 135
1825 - Portunus hastatus (Linnaeus) in Latreille; p. 189
1933 - Neptunus hastatus (Linnaeus) in de Miranda; p. 49
1936 - Lupa bastata (Linnaeus) in Nobre; p. 41, PI. 15, Text-
figs. 37-38
1940 - Neptunus hastatus (Linnaeus) in Bouvier; p. 249, Text-fig. 160,
PI. 9 (fig. 14)
1946 - Neptunus hastatus (Linnaeus) in Zariquiey Alvarez; p. 158,
PI. 15 (b-c)
1950 - Lupa bastata (Linnaeus) in Comaschi Caria; p. 324, PI. 1
1952 - Neptunus hastatus (Linnaeus) in Zariquiey Alvarez; p. 36
1956 - Neptunus hastatus (Linnaeus) in Monod; p. 203, Text-
figs. 232-235
1956 -Lupa bastata (Linnaeus) in Comaschi Caria; p. 288
1958 - Portunus hastatus (Linnaeus) in Holthuis & Gottlieb; p. 91
1968 - Portunus hastatus (Linnaeus) in Zariquiey Alvarez; p. 384,
Text-figs. 125 d-e, 126 c, 128 a-bl
1992 - Portunus hastatus (Linnaeus) in Falciai & Minervini; p. 20 1 -
202
Holotype: unknown.
Stratigraphic range: middle Miocene (Helvetian).
Occurrence: Sardegna.
Material: Comaschi Caria (1950) reported one
specimen from Bosa (Nuoro) (repository and catalogue
number unknown).
Portunus incertus (A. Milne Edwards, 1 860)
1860 - Neptunus incertus A. Milne Edwards; p. 244, PI. 5 (fig. 3)
191 Oa -Neptunus incertus A. Milne Edwards in Fabiani; p. 23
1915 - Neptunus incertus A.. Milne Edwards in Fabiani; p. 285
1929 - Neptunus incertus A. Milne Edwards in Glaessner; p. 268
1996 - Portunus incertus (A. Milne Edwards) in Beschin, Checchi &
Ungaro; p. 16
2001- Portunus ( Portunus ) incertus (A. Milne Edwards) in De Angeli
& Beschin; p. 30
Holotype: unknown.
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: A. Milne Edwards (1860) reported this spe-
cies from Salcedo (Vicenza) (repository and catalogue
number unknown).
Portunus kochi (Bittner, 1893)
1893 - Neptunus Kochii Bittner; p. 22, PI. 1 (figs. 1, 1 a)
1929 - Neptunus Kochi Bittner in Glaessner; p. 268
1929 - Neptunus kochi Bittner in Lorenthey & Beurlen; p. 185, PI. 13
(figs. 5 a-c, 8 a-b)
1969 - Neptunus kochi Bittner in Via Boada; p. 216
1996 - Portunus kochi (Bittner) in Beschin, Checchi & Ungaro; p. 14,
Text-fig. 3, PI. 1 (figs. 1-3)
2001 - Portunus ( Portunus ) kochi (Bittner) in De Angeli & Beschin;
p. 30, Text-fig. 25
Holotype: unknown.
Stratigraphic range: Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (1996) reported three speci¬
mens (MCZ from 1564 to 1566) from Castelgomberto
(Vicenza).
Note: this species is also reported from thè middle
Eocene of Hungary.
Portunus larteti (A. Milne Edwards, 1860)
1860 - Neptunus Larteti A. Milne Edwards; p. 237, PI. 5 (figs. 2,
2 b)
1875 - Neptunus Lartetii A. Milne Edwards in Bittner; p. 80
1 9 1 Oa - Neptunus Larteti A. Milne Edwards in Fabiani; p. 22
1915 - Neptunus Larteti A. Milne Edwards in Fabiani; p. 285
1929 - Neptunus larteti A. Milne Edwards in Glaessner; p. 268
1996 - Portunus larteti (A. Milne Edwards) in Beschin, Checchi &
Ungaro; p. 16
2001 - Portunus ( Portunus ) larteti (A. Milne Edwards) in De Angeli
& Beschin; p. 30
Holotype: unknown
Stratigraphic range: Tertiary.
Occurrence: Veneto.
Material: A. Milne Edwards (1860) reported this spe¬
cies from thè “nummulitico” of Vicenza (repository and
catalogue number unknown).
Portunus monspeliensis (A. Milne Edwards, 1860)
1860 - Neptunus monspeliensis A. Milne Edwards; p. 106, PI. 4
(fig. 1), PI. 5 (fig. 1)
1860 - Neptunus granulatus A. Milne Edwards; p. 115, PI. 3 (fig. 1),
PI. 7 (fig. 2), nov. syn.
1888 - Neptunus granulatus A. Milne Edwards in Ristori; p. 215,
PI. 4 (figs. 5-11)
1893 - Neptunus granulatus A. Milne Edwards in Bittner; p. 1 1
1897 - Neptunus monspeliensis A. Milne Edwards in Roman; p. 128
60
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1898 - Neptunus granulatus A. Milne Edwards in Lòrenthey; p. 92,
PI. 9 (figs. 2-3)
1901 - Neptunus granulatus A. Milne Edwards in Blanckenhorn;
p. 76, 112
1909 - Neptunus granulatus A. Milne Edwards in Lòrenthey; p. 242,
PI. 2 (figs. 1-2)
191 1 - Neptunus granulatus A. Milne Edwards in Toula; p. 48, Text-
figs. 1-2
1927 - Neptunus granulatus A. Milne Edwards in Van Straelen; p. 86
1 928 - Neptunus granulatus A. Milne Edwards in Glaessner; p. 1 83
1929 - Neptunus granulatus A. Milne Edwards in Glaessner; p. 267
1 929 - Neptunus granulatus A. Milne Edwards in Lòrenthey & Beur-
len; p. 188, PI. 13 (figs. 3-4), PI. 14 (figs. 1-4)
1 950-Neptunus granulatus A. Milne Edwards in Comaschi Caria; p. 326
1956 - Neptunus granulatus A. Milne Edwards in Comaschi Caria;
p. 288, PI. 1 (figs. 1-7), PI. 2 (figs. 1-6), PI. 3 (figs. 1-2)
1962 - Neptunus granulatus A. Milne Edwards in Zbyszewski & Da
Veiga Ferreira; p. 286
1964-65 - Neptunus granulatus A. Milne Edwards in Da Veiga Fer¬
reira; p. 150
1965 - Neptunus granulatus A. Milne Edwards in Varola; p. 296
1968 - Neptunus cfr. N. granulatus A. Milne Edwards in Stancu &
Andreescu; p. 466, PI. 7 (fig. 85)
1979 - Portunus monspeliensis (A. Milne Edwards) in Muller; p. 274,
280,288, PI. 18
1979 - Portunus granulatus A. Milne Edwards in Forster; p. 94
1984 - Portunus monspeliensis (A. Milne Edwards) in Muller; p. 79,
PI. 62 (figs. 1-2)
1987 - Portunus monspeliensis (A. Milne Edwards) in Allasinaz;
p. 539, PI. 4 (figs. 1-2)
1991 - Portunus monspeliensis (A. Milne Edwards) in Marras & Ven¬
tura; p. 108, PI. 1, PI. 2 (figs. 1-4), PI. 3 (figs. 1-3)
1998 - Portunus monspeliensis (A. Milne Edwards) in Muller; p. 29
2006a - Portunus monspeliensis (A. Milne Edwards) in De Angeli &
Garassino; p. 283, Text-fig. 8
Holotype: MNHN R03774.
Stratigraphic range: lower Miocene (Burdigalian) -
middle Miocene (Langhian).
Occurrence: Piemonte, Friuli-Venezia Giulia, Emilia
Romagna, Puglia, Sardegna.
Material: Ristori (1888) reported many specimens
from Monte S. Michele, Flussio and Tresnuraghes
(Cagliari), housed in Lovisato’s collection (repository and
catalogue number unknown), some specimens from Lecce
and one specimen from S. Maria Vigliana (Bologna),
today lost; Lòrenthey (1909) reported some specimens
from Cadreas, Portotorres, Sedin, Ardara (Sassari) and
Planargia (Cagliari) (repository and catalogue number
unknown); Comaschi Caria (1956) reported one specimen
from Bonorva and Bosa (Nuoro) (repository and cata¬
logue number unknown); Allasinaz (1987) reported three
specimens (MCSN 4M, 91M, 222) from Fontanino di Rio
Caramagna, Ciglione, and Ponzone (Acqui); Marras &
Ventura (1991) reported some specimens from Sassari
(repository and catalogue number unknown); De Angeli
& Garassino (2006a) reported two specimens (MSNM
Ì26584, Ì26585) from Meduno (Pordenone).
Note: this species is also reported from thè Miocene of
Hungary, Spain, Austria, France, and Egypt.
Portunus neogenicus Muller, 1979
1979 - Portunus neogenicus Muller; p. 280, PI. 19
1984 - Portunus neogenicus Muller in Muller; p. 80, PI. 62 (figs. 3-4)
1991 - Portunus neogenicus Miiller in Marras & Ventura; p. 110,
PI. 3 (fig. 4)
Holotype: (MAFI).
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Marras & Ventura (1991) reported one
specimen from Sassari (repository and catalogue number
unknown).
Note: this species is also reported in thè Miocene of
Hungary.
Portunus cfr. P. radobojanus (Bittner, 1884)
1 892c - Neptunus affi radobojanus Bittner in Ristori; p. 161
191 Oa - Neptunus sp. (N. aff. Radobojanus Bittner) in Fabiani; p. 23
1929 - Neptunus cfr. radobojanus Bittner in Glaessner; p. 269
1996 - Portunus aff. radoboyanus (Bittner) in Beschin, Checchi &
Ungaro; p. 16
2001 - Portunus ( Portunus ) aff. radobojanus (Bittner) in De Angeli &
Beschin; p. 31
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: Ristori (1892c) reported four or five speci¬
mens from Chiavon (Vicenza) (repository and catalogue
number unknown).
Portunus cfr. P. stenaspis (Bittner, 1884)
1892c -Neptunus aff. stenaspis Bittner in Ristori; p. 161
1 9 1 Oa - Neptunus sp. ( N . aff. stenaspis Bittner) in Fabiani; p. 23
1929 - Neptunus cf. stenaspis Bittner in Glaessner; p. 269
1996 - Portunus aff. stenaspis Bittner in Beschin, Checchi & Ungaro;
p. 16
2001 - Portunus ( Portunus ) aff. stenaspis (Bittner) in De Angeli &
Beschin; p. 31
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: Ristori (1892c) reported one specimen from
Chiavon (Vicenza) (repository and catalogue number
unknown).
Portunus suessi (Bittner, 1875)
1875 - Neptunus Suessi Bittner; p. 80, PI. 4 (fig. 1 a-d)
191 Oa -Neptunus Suessi Bittner in Fabiani; p. 23
1915 - Neptunus Suessi Bittner in Fabiani; p. 285
1929 - Neptunus Suessi Bittner in Glaessner; p. 297
2001 - Portunus ( Portunus ) suessi (Bittner) in De Angeli & Beschin;
p. 31
Holotype: unknown
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Bittner (1875) reported this species from
Laverda (Vicenza) (repository and catalogue number
unknown).
Portunus tuberculatus Roux, 1828
1828 - Portunus tuberculatus Roux; PI. 32 (figs. 1-5)
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
61
1914 - Portunus tuberculatus Roux in M. Gemmellaro; p. 88, PI. 1
(figs. 22-23)
1929 - Portunus tuberculatus Roux in Glaessner; p. 336
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported one speci¬
men from Ficarazzi (Palermo) (MGUP, catalogue number
unknown).
Portunus vicentinus (A. Milne Edwards, 1 860)
1860 - Neptunus Vicentinus A. Milne Edwards; p. 238, PI. 6
(fig. 1 a-b)
1875 - Neptunus Vicentinus A. Milne Edwards in Bittner; p. 80
1 9 1 Oa - Neptunus vicentinus A. Milne Edwards in Fabiani; p. 23
1929 - Neptunus vicentinus A. Milne Edwards in Glaessner;
p. 269
1996 - Potunus vicentinus (A. Milne Edwards) in Beschin, Checchi &
Ungaro; p. 16
2001 - Portunus ( Portunus ) vicentinus (A. Milne Edwards) in
De Angeli & Beschin; p. 31
Holotype: MNHN R03818.
Stratigraphic range: Tertiary.
Occurrence: Veneto.
Material: A. Milne Edwards (1860) reported one
specimen (MNHN R03818) from thè “nummulitico” of
Vicenza.
Portunus sp.
1892c - Neptunus sp. in Ristori; p. 161
1929 - Neptunus sp. in Glaessner; p. 270
Stratigraphic range: Oligocene.
Occurrence: Veneto.
Material: Ristori (1892c) reported some specimens
fforn Chiavon (Vicenza) (repository and catalogue
number unknown).
Portunus sp.
1 895 - Portunus sp. in Crema; p. 677
1929 - Portunus sp. in Glaessner; p. 336
Stratigraphic range: upper Miocene (Tortonian) -
upper Pliocene (Piacentian).
Occurrence: Piemonte.
Material: Crema (1895) reported some specimens
from Torino and Villavernia (Alessandria), today
lost.
Portunus sp.
1909 - Neptunus sp. in Lòrenthey; p. 245
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported some specimens
from Monte S. Michele (Cagliari) (repository and cata¬
logue number unknown).
Portunus sp.
1975 - Portunus sp. in Secretan; p. 358, PI. 3
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported one specimen
(MSNV M3) from Monte Bolca (Verona).
Portunus ( Achelous ) De Haan, 1833
Type species: Portunus spinimanus Latreille, 1819.
Stratigraphic range: Oligocene - Recent.
Portunus ( Achelous ) obtusus A. Milne Edwards, 1860
1860 -Achelous obtusus A. Milne Edwards; p. 245, PI. 3 (fig. 2)
1875 - Achelous obtusus A. Milne Edwards in Bittner; p. 83
1910a -Achelous obtusus A. Milne Edwards in Fabiani; p. 23
1915 - Achelous obtusus A. Milne Edwards in Fabiani; p. 285
1929 - Achelous obtusus A. Milne Edwards in Glaessner; p. 50
2001 - Portunus ( Achelous ) obtusus A. Milne Edwards in De Angeli
& Beschin; p. 32
Holotype: MNHN R03807.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: A. Milne Edwards (1860) reported
one specimen (MNHN R03807) from Salcedo (Vi¬
cenza).
Rakosia Miiller, 1984
Type species: Rakosia carupoìdes Miiller, 1984.
Stratigraphic range: lower Oligocene (Rupelian) -
Miocene.
Rakosia grumiensis Beschin, De Angeli & Checchi, 2001
2001 - Rakosia grumiensis Beschin, De Angeli & Checchi; p. 23,
Text-fig. 7, PI. 2 (figs. 3, 6)
2001 - Rakosia grumiensis Beschin, De Angeli & Checchi in De Angeli
& Beschin; p. 3 1
Holotype: MCZ 2128 (EG. 286483).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported 3 1 specimens
(MCZ from 2128 to 2150, from 2156 to 2163) from Ca-
stelgomberto (Vicenza).
Scylla De Haan, 1833
Type species: Cancer serratus Forskàl, 1775.
Stratigraphic range: Eocene - Recent.
Scylla sp.
1892 - Scylla sp. in Ristori; p. 163
1910 a -Scylla sp. in Fabiani; p. 23
1929 - Scylla sp. in Glaessner; p. 375
2001 - Scylla sp. in De Angeli & Beschin; p. 32
Stratigraphic range: lower Oligocene (Rupelian).
62
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Occurrence: Veneto.
Material: Ristori (1892c) reported two specimens
from Chiavon (Vicenza) (repository and catalogue
number unknown).
Indeterminates
Genus et species indeterminate
1998b - Genus et species indeterminate in Bravi & Garassino; p. 112,
Text-fig. 15
Stratigraphic range: Lower Cretaceous (Albian).
Occurrence: Campania.
Material: Bravi & Garassino (1998b) reported one
specimen (M 21840) from Petina (Salerno).
Superfamily Xanthoidea MacLeay, 1838
Family Carpiliidae Ortmann, 1893
Palaeocarpilius A. Milne Edwards, 1862
Type species: Cancer macrochelus Desmarest, 1822.
Stratigraphic range: Eocene - Miocene.
Palaeocarpilius aquitanicus A. Milne Edwards, 1 862
1 847 - Cancer Bosciì in Burguet; p. 280
1850 - Cancer Boscii in d’ Archiac; p. 448
1861 - Cancer Boscii in Michelotti; p. 139
1862 - Palaeocarpilius aquitanicus A. Milne Edwards; p. 57, PI. 4
(figs. 4, 4 a)
1875 - Palaeocarpilius platycheilus (Reuss) in Bittner; p. 84, PI. 3
(fig. 4)
1 889 - Palaeocarpilius macrocheilus (Desmarest) in Ristori; p. 398
1929 - Palaeocarpilius aquitanicus A. Milne Edwards in Glaessner;
p. 292
1974 - Palaeocarpilius macrocheilus (Desmarest) in Mastrorilli; p. 4
1987 - Palaeocarpilius macrocheilus (Desmarest) in Allasinaz; p. 541,
PI. 5 (figs. 1-2)
1996 - Palaeocarpilius macrochelus (Desmarest) in Beschin, Checchi
& Ungaro; p. 16, Text-fig. 4, PI. 2 (figs. 1-2)
2001 - Palaeocarpilius macrochelus (Desmarest) in Beschin,
De Angeli & Checchi; p. 24, PI. 3 (fig. 1 a-b)
2003 - Palaeocarpilius aquitanicus A. Milne Edwards in Schweitzer;
p. 1112
Holotype: MNHN R03772 (syntype, cast).
Stratigraphic range: Oligocene - Miocene.
Occurrence: Piemonte, Veneto.
Material: Ristori (1889) reported many specimens
(P. macrocheilus ) from Sassello (Savona), housed in
don Perrando’s collection today in “DIP.TE.RIS” (we
identified two originai specimens: 2388/Sa-II-S48 (ex
Sa-II-S 1 84) and 2390/Sa-II-S 1 87 (ex Sa-II-S184), not
figured by thè author); Allasinaz (1987) reported one
specimen (P. macrocheilus ) from Ciglione (Ovada)
(MCSN, catalogue number unknown); Beschin et al.
(1996) reported tour specimens (P. macrochelus , MCZ
1234, 1568, 1569, 1570) from Castelgomberto (Vicen¬
za); Beschin et al. (2001) reported three specimens
(P. macrochelus, MCZ form 2152 to 2154) from Ca¬
stelgomberto (Vicenza).
Note: this species reported from thè lower Oligocene
(Rupelian) of N Italy, was usually confused with P. mac¬
rochelus Desmarest, 1822, which has a carapace with
eight rounded lobes and widespread in thè levels of thè
upper Eocene (Beschin et al., 2006). This species is also
reported from thè lower Oligocene of SW France.
Palaeocarpilius macrochelus (Desmarest, 1 822)
Cancer lapidescens Rumphius; p. 84, PI. 61 (fig. 3)
1622 - Cancer lapidescens Rumphius in Museum Calceolarianum
Veronense; p. 407
Cancer lapidescens Rumphius in Aldovrandi (Museum Metal-
licum); p. 461
1656 - Cancer lapidescens Rumphius in Museo di Moscardi; p. 179
1822 - Cancer macrochelus Desmarest; p. 91, PI. 7 (figs. 1-2)
1 822 - Cancer Boscii Desmarest; p. 94, PI. 8 (figs. 3-4)
1 822 - Brachyurites antiquus Schlotheim; p. 26, PI. 1
Cancer antiquus Quenstedt; p. 26, Text-fig. 1
Carpilius macrochelus A. Milne Edwards; p. 380, PI. 1
1834 - Cancer boscii Desmarest in A. Milne Edwards; p. 379, PI. 1,
nov. syn.
1838 - Cancer macrochelus Desmarest in Mantell; p. 218, PI. 25
1854 - Cancer boscii Desmarest in Catullo; p. 1
1859 - Atergatis boscii (Desmarest) in Reuss; p. 30, PI. 9 (figs. 4-9),
PI. 10 (fig. 1), PI. 11 (figs. 1-4), PI. 12 (figs. 1-2)
1859 - Atergatis stenura Reuss; p. 30, PI. 11 (figs. 5-7), nov. syn.
1859 - Atergatis platycheilus Reuss; p. 36, PI. 10 (figs. 2-3), nov. syn.
1862 - Palaeocarpilius macrocheilus (Desmarest) in A. Milne
Edwards; p. 186, PI. 1 (fig. 2), PI. 2 (fig. 1), PI. 3 (fig. 1)
1867 - Artergatis boscii (Desmarest) in Fraas; p. 300
1875 - Palaeocarpilius macrocheilus (Desmarest) in Bittner; p. 83
1 875 - Palaeocarpilius stenurus (Reuss) in Bittner; p. 84
1883 - Palaeocarpilius macrocheilus (Desmarest) in Bittner; p. 311
1 883 - Palaeocarpilius platycheilus (Reuss) in Bittner; p. 3 1 1
1885 - Palaeocarpilius macrocheilus (Desmarest) in Noetling; p. 487,
489, PI. 4 (fig. 2)
1886 - Palaeocarpilius macrocheilus var. coronata Bittner; p. 44,
PI. 1 (fig- 1)
1893 - Palaeocarpilius macrocheilus (Desmarest) in Bittner; p. 20
1 895 - Cancer ( Palaeocarpilius ) macrochelus (Desmarest) in De Gre¬
gorio; p. 13, PI. 4 (figs. 1-5)
1895 - Harpactocarcinus supragigas De Gregorio; p. 13, PI. 6
(figs. 1-3), nov. syn.
1896 - Palaeocarpilius macrocheilus (Desmarest) in Vinassa
de Regny; p. 127, PI. 2 (fig. 2 a-b)
1898 - Palaeocarpilius macrocheilus (Desmarest) in Lòrenthey; p. 36
1899 - Palaeocarpilius macrocheilus (Desmarest) in Oppenheim;
p. 59
1901 - Palaeocarpilius macrocheilus (Desmarest) in Oppenheim;
p. 281
1908 - Palaeoocarpilius macrocheilus (Desmarest) in Fabiani; p. 210,
236
1909 - Palaeocarpilius macrocheilus (Desmarest) in Lòrenthey;
p. 132
191 Oa — Palaeocarpilius macrocheilus (Desmarest) in Fabiani; p. 24
1915 - Palaeocarpilius macrocheilus (Desmarest) in Fabiani; p. 284,
285
1915 - Palaeocarpilius macrocheilus (Desmarest) in Dainelli; p. 699
1929 - Palaeocarpilius macrocheilus (Desmarest) in Glaessner; p. 292
1 944 - Palaeocarpilius macrocheilus (Desmarest) in Checchia-Rispoli;
p. 109
1946 - Palaeocarpilius macrocheilus (Desmarest) in Stubblefield;
p. 513, PI. 8 (figs. 2-6)
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
63
1953 - Palaeocarpilius macrocheilus (Desmarest) in Roger; p. 359,
PI- 2 (fig. 1)
1962 - Palaeocarpilius macrocheilus (Desmarest) in Piccoli & Mocel-
lini; p. 38, 48, 78
1969 - Palaeocarpilius macrochelus (Desmarest) in Glaessner;
p. 520, Text-fig. 328 (1)
1969 - Palaeocarpilius macrocheilus (Desmarest) in Via Boada;
p. 403
1995 - Palaeocarpilius macrochelus (Desmarest) in De Angeli; p. 1 6
2001 - Palaeocarpilius macrochelus (Desmarest) in De Angeli &
Beschin; p. 32, Text-fig. 27
2003 - Palaeocarpilius macrocheilus (Desmarest) in Schweitzer;
p. 1112
2004 - Palaeocarpilius macrochelus (Desmarest) in Beschin &
De Angeli; p. 20
2006 - Palaeocarpilius macrochelus (Desmarest) in Beschin,
De Angeli, Checchi & Mietto; p. 107, PI. 3 (figs. 3, 4 a-b)
Holotype: MNHN R03830.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Desmarest (1822) reported one specimen
(MNHN R03830) from China?, but probably discovered
in thè “nummulitico” of Verona; A. Milne Edwards ( 1 862-
1 865) reported four specimens (MNHN R038 14, R03829,
R03830, A24548) without pointing out thè fossiliferous
localities, probably discovered in thè “nummulitico” of
Verona; Vinassa de Regny (1896) reported one specimen
from Avesa (Verona) (repository and catalogue number
unknown); Reuss (1859), Bittner (1886), and Fabiani
(1910a) reported this specimens from some localities of
Veneto (repository and catalogue number unknown); De
Gregorio (1895) reported three specimens from Valrovina
(Vicenza) (repository and catalogue number unknown),
and one specimen from Verona (MGV, catalogue number
unknown); Dainelli (1915) reported two specimens from
Monte Plauris (Udine) (repository and catalogue number
unknown); Piccoli & Mocellin (1962) reported one speci¬
men from Priabona-Monte di Malo (Vicenza) (MGPD,
catalogue number unknown); De Angeli (1995) reported
one specimen from “Fontanella” di Grancona (Vicenza)
(MCZ, catalogue number unknown); Beschin et al.
(2006) reported fìve specimens (MCZ 1191, 1235, 1327,
2445, 1446) from Grotta della Poscola, Buso della Rana,
and Val Segato (Vicenza).
Note: thè study of some carapaces from thè Oli¬
gocene of Vicenza area and Bacino Ligure-Piemontese
has pointed out thè presence of seven anterolateral lobes
instead of eight, as reported for thè specimens of P. ma¬
crochelus, discovered in thè levels of thè upper Eocene.
The presence of seven alterolateral lobes is typical of
P. aquitanicus A. Milne Edwards, 1862, reported from
thè levels of thè lower Oligocene (Rupelian) of France.
Palaeocarpilius macrochelus is also reported from thè
Eocene of Hungary, France, Egypt, and Somalia.
Palaeocarpilus simplex Stoliczka, 1 87 1
1871 - Palaeocarpilius simplex Stoliczka; p. 1 1, PI. 5 (fig. 6)
1 875 - Palaeocarpilius anodon Bittner; p. 85, PI. 2 (fig. 3), nov. syn.
1909 - Palaeocarpilius simplex Stoliczka in Lòrenthey; p. 127, PI. 1
(figs. 3-4)
191 Oa — Palaeocarpilius anodon Bittner in Fabiani; p. 24
1915 - Palaeocarpilius anodon Bittner in Fabiani; p. 284
1929 - Palaeocarpilius simplex Stoliczka in Glaessner; p. 294
1933 - Palaeocarpilius anodon Bittner in Di Salvo; p. 18, PI. 1
(fig. 1 a-c)
1959 - Palaeocarpilius simplex Stoliczka in Via Boada; p. 380
1969 - Palaeocarpilius simplex Stoliczka in Via Boada; p. 231, PI. 23
(figs. 3-4)
1982 - Palaeocarpilius anodon Bittner in Busulini, Tessier & Visentin;
p. 81
1994 - Palaeocarpilius simplex Stoliczka in Beschin, Busulini,
De Angeli & Tessier; p. 1 87, PI. 9 (fig. 1 a-b)
2001 - Palaeocarpilius simplex Stoliczka in De Angeli & Beschin;
p. 33
2003 - Palaeocarpilius simplex Stoliczka in Schweitzer; p. 1112
2003 - Palaeocarpilius anodon Bittner in Schweitzer; p. 1112
2004 - Palaeocarpilius simplex Stoliczka in Beschin, Busulini, De
Angeli & Tessier; p. 115
2004 - Palaeocarpilius simplex Stoliczka in Beschin & De Angeli; p. 20
2005 - Palaeocarpilius simplex Stoliczka in Beschin, De Angeli, Chec¬
chi & Zarantonello; p. 23, PI. 4 (fig. 9)
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto, Sicilia.
Material: Bittner (1875) reported one specimen
from S. Giovanni Ilarione (Verona) (type of P. anodon
Bittner, 1875 = MNHB K.lll, MB. A. 661); Di Salvo
(1933) reported one specimen from Ponte di Castronuovo
(Palermo) housed in Di Salvo’s collection (repository
and catalogue number unknown); Busulini et al. (1982)
reported six specimens from “Main” quarry of Arzignano
(Vicenza) (repository and catalogue number unknown);
Beschin et al. (1994) reported one specimen (MCZ 1205)
from “Boschetto” quarry of Nogarole Vicentino (Vice¬
nza); Beschin et al. (2005) reported fìve specimens (MCZ
2302, 2303, 2324; MV 48, 04/17) from Grola di Comedo
Vicentino (Vicenza).
Note: this species is also reported from thè Eocene of
Pakistan, Egypt, and Spain.
Palaeocarpilius valrovinensis (De Gregorio, 1895)
1895 - Cancer (Harpactocarcinus) Valrovinensis De Gregorio; p. 12,
PI. 5 (figs. 1-4)
191 Oa - Harpactocarcinus valrovinensis De Gregorio in Fabiani; p. 26
1915 - Harpactocarcinus valrovinensis De Gregorio in Fabiani; p. 285
1929 - Harpactocarcinus valrovinensis De Gregorio in Glaessner;
p. 206
2003 - Palaeocarpilius valrovinensis (De Gregorio) in Schweitzer;
p. 1119
Holotype: unknown.
Stratigraphic range: Eocene (Lutetian).
Occurrence: Veneto.
Material: De Gregorio (1895) reported one speci¬
men from Valrovina (Vicenza) (repository and catalogue
number unknown).
Family Domeciidae Ortmann, 1893
Jonesius Sankarankutty, 1 962
Type species: Jonesius minuta Sankarankutty, 1962.
64
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: Oligocene - Recent.
Jonesius oligocenicus (Beschin, De Angeli & Checchi,
2001)
2001 - Maldivia oligocenica Beschin, De Angeli & Checchi; p. 24,
Text-fig. 8, PI. 3 (fig. 2)
2001 - Maldivia oligocenica Beschin, De Angeli & Checchi in
De Angeli & Beschin; p. 37
2004 - Maldivia oligocenica Beschin, De Angeli & Checchi in Beschin
& De Angeli; p. 21
2005 - Jonesius oligocenicus (Beschin, De Angeli, Checchi) in
Schweitzer; p. 626
Holotype: MCZ 2121 (I.G. 286476).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported three speci-
mens (MCZ 2121, 2164, 2165) from Castelgomberto
(Vicenza).
Family Goneplacidae MacLeay, 1838
Brancìtioplax Rathbun, 1916
Type species: Branchioplax was hingtoniana Rathbun,
1916.
Stratigraphic range: Eocene.
Branchioplax albertìi De Angeli & Beschin, 2002
2002 - Branchioplax albertìi De Angeli & Beschin; p. 126, Text-
figs. 2-3
2004 - Branchioplax albertii De Angeli & Beschin in Beschin &
De Angeli; p. 21
2006 - Branchioplax albertii De Angeli & Beschin in Beschin,
De Angeli, Checchi & Mietto; p. 108, PI. 3 (figs. 1-2)
Holotype: MCZ 2062 (I.G. 286340).
Stratigraphic range: middle Eocene (Lutetian) -
upper Eocene (Priabonian).
Occurrence: Veneto.
Material: De Angeli & Beschin (2002) reported two
specimens (MCZ 2062, 2063) from “Main” quarry of
Arzignano (Vicenza); Beschin et al. (2006) reported four
specimens (MCZ 1328, from 2447 to 2449) from Buso
della Rana (Vicenza).
Chlinocephalus Ristori, 1886
Type species: Chlinocephalus demissifrons Ristori,
1886.
Stratigraphic range: Pliocene.
Chlinocephalus demissifrons Ristori, 1 886
1886 - Chlinocephalus demissifrons Ristori; p. 101, PI. 2 (figs. 5-6)
189 la - Titanocarcinus sculptus Ristori; p. 6, PI. 1 (figs. 1-2), nov.
syn.
1929 - Chlinocephalus demissifrons Ristori in Glaessner; p. 1 13
1981 - Chlinocephalus demissifrons Ristori in Delle Cave; p. 46
2004 - Chlinocephalus demissifrons Ristori in Garassino, De Angeli,
Gallo & Pasini; p. 275, Text-figs. 15-16
Syntype: IGF 628E (figured by Ristori in PI. 2, figs.
5-6).
Stratigraphic range: Pliocene.
Occurrence: Liguria, Piemonte.
Material: Ristori (1886) reported one specimen
from Fornaci (Savona) today lost; Garassino et al.
(2004) reported one specimen (PU 41 187) from Cossato
(Biella).
Note: Garassino et al. (2004) pointed out that
thè assignment of Titanocarcinus sculptus Ristori,
1891, from thè Pliocene clays of Mucigliani (Siena
- Toscana) to Titanocarcinus A. Milne Edwards, 1863,
is uncertain. In fact, thè shape of thè front, thè size
and location of thè orbits, thè presence of two spines
besides thè extraorbital spine on anterolateral mar-
gins, and thè transverse relief on thè cardiac region
are present also in Chlinocephalus demissifrons. The
specimen of T. sculptus studied by Ristori (189 la)
could be a juvenile stage (maximum width of carapace
= 2 mm; maximum length of carapace = 1.5 mm) of
Chlinocephalus demissifrons. The specimen studied by
Ristori ( 1 89 1 a) is housed in thè Museo dei Fisiocritici
di Siena (MFS 4624).
Corallicarcinus Miiller & Collins, 1991
Type species: Neptocarcinus spinosus Lòrenthey,
1929.
Stratigraphic range: upper Eocene - lower Oligo¬
cene.
Corallicarcinus sp.
2001 - Corallicarcinus sp. in Beschin, De Angeli & Checchi; p. 26,
PI. 3 (fig. 3)
2001 - Corallicarcinus sp. in De Angeli & Beschin; p. 37
2004 - Corallicarcinus sp. in Beschin & De Angeli; p. 21
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported one specimen
(MCZ 2127) from Castelgomberto (Vicenza).
Gollincarcinus Beschin & De Angeli, 2004
Type species: Gollincarcinus levis Beschin & De
Angeli, 2004.
Stratigraphic range: middle Eocene (Lutetian).
Gollincarcinus levis Beschin & De Angeli, 2004
2004 - Gollincarcinus levis Beschin & De Angeli; p. 15, Text-fig. 2,
PI. 1 (figs. 1-2, 3 a-e)
2005 - Gollincarcinus levis Beschin & De Angeli in Beschin,
De Angeli, Checchi & Zarantonello; p. 26, PI. 5 (fig. 4 a-b)
Holotype: MCZ 2405 (I.G. 305 1 13).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin & De Angeli (2004) reported five
specimens (MCZ 2305, 2405, 2406, 2407, 2408) from
“Main” quarry of Arzignano (Vicenza), “Boschetto”
quarry of Nogarole Vicentino (Vicenza), and Grola di
Comedo Vicentino (Vicenza); Beschin et al. (2005)
CATALOG AND BIBLIOGRAPHY OF THE FOSS1L STOMATOPODA AND DECAPODA FROM ITALY
65
reported one specimen (MCZ 2305) from Grola di
Comedo Vicentino (Vicenza).
Goneplax Leach, 1814
Type species: Cancer rhomboides Linnaeus, 1758.
Stratigraphic range: Miocene - Recent.
Goneplax gulderi Bachmayer, 1953
1895 - Goneplax sacci Crema; p. 674, Text-fig. 15, nov. syn.
1907 - Goneplax cfr. sacci Crema in Lorenthey; p. 94, PI. 3 (figs. 4 a-c,
6-7), PI. 4 (fìg. 6)
1909 - Goneplax cfr. sacci Crema in Lorenthey; p. 249, PI. 1 (fig. 4 a-c,
6-7), PI. 2 (fig. 6)
1929 - Goneplax saccoi Crema in Glaessner; p. 199
1948 - Goneplax cfr. saccoi Crema in Via Boada; p. 146, PI. 1 (fig. 6)
1953 - Goneplax gulderi Bachmayer; p. 143, PI. 9 (figs. 1-3)
1984 - Goneplax gulderi Bachmayer in Miiller; p. 96, PI. 93
(figs. 2-3)
1988 - Goneplax cf. saccoi Crema in Solé & Via Boada; p. 34
1993 - Goneplax gulderi Bachmayer in Miiller; p. 23, Text-fig. 1 1 K
1998 - Goneplax gulderi Bachmayer in Mayoral, Miiller & Muniz;
p. 508, Text-fig. 2 (5)
1998 - Goneplax gulderi Bachmayer in Miiller; p. 38
2004a - Goneplax gulderi Bachmayer in Garassino & De Angeli;
p. 45
Holotype: unknown.
Stratigraphic range: Miocene - Pliocene.
Occurrence: Piemonte, Sardegna.
Material: Crema (1895) reported one specimen from
Monte Capriolo (Bra - Cuneo), today lost; Lorenthey
(1909) reported some specimens from Capo S. Marco
(Cagliari), housed in Lovisato’ collection (repository and
catalogue number unknown).
Note: Miiller (1993) pointed out that Goneplax sacci
Crema, 1895, and G. cfr. G. sacci described by Lorenthey
(1907, 1909) are synonyms of G. gulderi Bachmayer,
1953. This species is also reported from thè Miocene of
Austria, Spain and Hungary.
Goneplax rhomboides (Linnaeus, 1758)
1758 - Cancer rhomboides Linnaeus; p. 626
1814 - Goneplax angolata (Bell) in Leach; p. 430
1 8 1 6 - Gonoplax bispinosa Leach; p. 420, nov. syn.
1822 - Goneplax impressa Desmarest; p. 102, PI. 8 (figs. 13-14), nov.
syn.
1861 - Goneplax rhomboides (Linnaeus) in A. Milne Edwards; p. 88
1863 - Gonoplax angulata (Bell) in Heller; p. 103
1863 - Gonoplax rhomboides (Linnaeus) in Heller; p. 104, PI. 3
(figs. 3-4)
1 886 - Goneplax impressa Desmarest in Ristori; p. 1 1 1
1886 - Goneplax formosa Ristori; p. 1 1 1, PI. 3 (figs. 11-13), nov. syn.
1886 - Goneplax meneghina Ristori; p. 114, PI. 3 (figs. 8-10), nov.
syn.
1 89 1 b — Gonoplax bispinosa Leach in Ristori; p. 20
1892a - Gonoplax meneghina Ristori in Ristori; p. 89
1896 - Gonoplax meneghina Ristori in Ristori; p. 506
1896 - Gonoplax rhomboides (Linnaeus) in Caullery; p. 405
1914 - Gonoplax cfr. rhomboides (Linnaeus) in M. Gemmellaro; p. 90,
PI. 1 (fig. 26)
1918 - Gonoplax angulata Heller in Pesta; p. 436, Text-fig. 144 a-b
1923 - Goneplax meneghini ? Ristori in Tettoni; p. 161
1929 - Goneplax rhomboides (Linnaeus) in Glaessner; p. 199
1929 - Goneplax formosa Ristori in Glaessner; p. 198
1929 - Goneplax meneghina Ristori in Glaessner; p. 199
1933 - Goneplax angulata (Bell) in de Miranda; p. 54
1936 - Goneplax angulata (Bell) in Nobre; p. 57, PI. 21 (fig. 40)
1940 - Goneplax angulata (Bell) in de Miranda; p. 29
1940 - Goneplax angulata (Bell) in Bouvier; p. 278, Text-fig. 176,
PI. 9 (fig. 2)
1946 - Goneplax angulata (Bell) in Zariquiey Alvarez; p. 162, PI. 18
(a-b)
1958 - Goneplax rhomboides (Linnaeus) in Holthuis & Gottlieb; p. 99
1959 - Goneplax rhomboides (Linnaeus) in Zariquiey Alvarez; p. 5
1961 - Goneplax rhomboides (Linnaeus) in Holthuis; p. 57
1961 - Goneplax angulata (Bell) in Nunes-Ruivo; p. 28
1968 - Goneplax rhomboides (Linnaeus) in Zariquiey Alvarez; p. 414,
Text-figs. 1 e, 138 a-b
1981 - Goneplax formosa (Linnaeus) in Delle Cave; p. 48
1992 - Goneplax rhomboides (Linnaeus) in Falciai & Minervini;
p. 238, PI. 27 (fìg.l)
2004a - Goneplax rhomboides (Linnaeus) in Garassino & De Angeli;
p. 44, Text-figs. 12-15
2004 - Goneplax rhomboides (Linnaeus) in Garassino, De Angeli,
Gallo & Pasini; p. 274, Text-fig. 14
Holotype: unknown.
Stratigraphic range: Pliocene - Pleistocene.
Occurrence: Piemonte, Emilia Romagna, Toscana,
Lazio, Sicilia.
Material: Desmarest ( 1 822) reported some specimens
from Monte Mario (Roma) (repository and catalogue
number unknown); A. Milne Edwards (1861) reported
some specimens from Palermo (repository and catalogue
number unknown); Ristori (1886) reported four speci¬
mens from Rapolano (Siena) (IGF, catalogue number
unknown); Ristori (1891b) reported one specimen from
Monte Mario (Roma), housed in Zuccari’s collection
(MGPUR, catalogue number unknown); M. Gemmellaro
(1914) reported one specimen from Ficarazzi (Palermo)
(MGUP, catalogue number unknown); Tettoni (1923)
reported some specimens from S. Venanzio (Modena)
(MGUM, catalogue number unknown); Garassino & De
Angeli (2004a) reported 13 specimens (MG 0566, from
0596 to 0600, 0601, 0602, 0612, 0614, 0621, 0632, 0633)
from Fiume Arda, Fiume Enza, and Torrente Stirane (Pia¬
cenza and Parma); Garassino et al. (2004) reported two
specimens (PU 41185, PU 41186) from Cocconato (Asti)
and Cossato (Biella).
Note: Miiller (1993) pointed out that Goneplax for¬
mosa Ristori, 1886 (holotype, IGF 612E, figured by Ri¬
stori in PI. 3, figs. 11-13) and G. meneghina Ristori, 1886
(syntypes, IGF 609E, figured by Ristori in PI. 3, fig. 10;
610E, figured by Ristori in PI. 3, figs. 8, 9, 9 a; 61 1 E not
figured) both from Rapolano (Siena) are synonyms of G.
rhomboides (Linnaeus, 1758). The specimens of these
species are housed in thè Museo di Geologia e Paleon¬
tologia dell’Università di Firenze. This species is also
reported from thè upper Pliocene of Great Britain.
Lessinioplax Beschin & De Angeli, 2004
Type species: Lessinioplax simplex Beschin & De
Angeli, 2004.
Stratigraphic range: middle Eocene (Lutetian).
66
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Lessinioplax rugosa Beschin & De Angeli, 2004
2004 - Lessinioplax rugosa Beschin & De Angeli; p. 20, Text-fìg. 4,
PI. 2 (figs. 2 a-b, 3)
Holotype: MCZ 1 1 75 (I.G. 2 1 1 690).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence; Veneto.
Material: Beschin & De Angeli (2004) reported two
specimens (MCZ 1175, 2414) from ”Albanello” quarry of
Nogarole Vicentino (Vicenza).
Lessinioplax simplex Beschin & De Angeli, 2004
2004 - Lessinioplax simplex Beschin & De Angeli; p. 18, Text-fìg. 3,
PI. 2 (figs. 1 a-c, 4 a-c)
Holotype: MCZ 2409 (I.G. 305117).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin & De Angeli (2004) reported five
specimens (MCZ 2409, 2410, 2411, 2412, 2413) from
“Main” quarry of Arzignano (Vicenza).
Magyarcarcinus Schweitzer & Karasawa, 2004
Type species: Palaeograpsus lóczyanus Lòrenthey,
1897.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Magyarcarcinus lóczyanus (Lòrenthey, 1 897)
1897 - Palaeograpsus lóczyanus Lòrenthey; p. 157, 168
1 898 - Palaeograpsus lóczyanus Lòrenthey in Lòrenthey; p. 69, PI. 4
(fig- 6)
1 929 - Palaeograpsus lóczyanus Lòrenthey in Glaessner; p. 296
1 929 - Palaeograpsus lóczyanus Lòrenthey in Lòrenthey & Beurlen;
p. 255, PI. 16 (fig. 1 a-e)
1994 - Palaeograpsus lóczyanus Lòrenthey in Beschin, Busulini,
De Angeli & Tessier; p. 196, PI. 1 1 (figs. 1-3)
1998 - Palaeograpsus lóczyanus Lòrenthey in Beschin, Busulini,
De Angeli, Tessier & Ungaro; p. 3 1 , figs. 1 5 (3), 1 6 (2)
2000 - Palaeograpsus lóczyanus Lòrenthey in Beschin, De Angeli &
Alberti; p. 15
2001 - Palaeograpsus lóczyanus Lòrenthey in De Angeli & Beschin;
p. 38, Text-fìg. 33
2001 - Carcinoplax lóczyanus (Lòrenthey) in Karasawa & Kato;
p. 272
2004 - Magyarcarcinus lóczyanus (Lòrenthey) in Schweitzer &
Karasawa; p. 76, Text-fig. 1 (3-5)
2004 - Magyarcarcinus lóczyanus (Lòrenthey) in Beschin &
De Angeli; p. 21
2005 - Magyarcarcinus lóczyanus (Lòrenthey) in Beschin, De Angeli,
Checchi & Zarantonello; p. 26
Holotype: MAFI E282, E283.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported six speci¬
mens (MCZ 1213, 1216, from 1417 to 1419, 1441) from
“Boschetto” quarry of Nogarole Vicentino (Vicenza);
Beschin et al. (1998) reported four specimens (MCZ
from 1517 to 1520) from “Rossi” quarry of Monte di
Malo (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Maingrapsus Tessier, Beschin, Busulini & De Angeli,
1999
Type species: Maingrapsus quadratus Tessier,
Beschin, Busulini & De Angeli, 1999.
Stratigraphic range: middle Eocene (Lutetian).
Maingrapsus quadratus Tessier, Beschin, Busulini &
De Angeli, 1999
1999 - Maingrapsus quadratus Tessier, Beschin, Busulini &
De Angeli; p. 98, Text-fig. 3, PI. 2 (figs. 1-3)
2001 - Maingrapsus quadratus Tessier, Beschin, Busulini & De Angeli
in De Angeli & Beschin; p. 38
2001 - Maingrapsus quadratus Tessier, Beschin, Busulini & De Angeli
in Karasawa & Kato; p. 272
2004 - Maingrapsus quadratus Tessier, Beschin, Busulini & De Angeli
in Beschin & De Angeli; p. 21
2004 - Maingrapsus quadratus Tessier, Beschin, Busulini & De Angeli
in Beschin, Busulini, De Angeli & Tessier; p. 116
Holotype: MCZ 1613 (I.G. 284680).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Tessier et al. (1999) reported five speci¬
mens (MCZ 1591, from 1601 to 1 603, 1613) from “Main”
quarry of Arzignano (Vicenza).
Note: Karasawa & Kato (2001) suggested thè ascrip-
tion of this genus to thè family Goneplacidae.
Paracorallicarcinus Tessier, Beschin, Busulini &
De Angeli, 1999
Type species: Paracorallicarcinus arcanus Tessier,
Beschin, Busulini & De Angeli, 1999.
Stratigraphic range: middle Eocene (Lutetian).
Paracorallicarcinus arcanus Tessier, Beschin, Busulini
& De Angeli, 1999
1999 - Paracorallicarcinus arcanus Tessier, Beschin, Busulini & De
Angeli; p. 95, Text-fig. 2, PI. 1 (figs. 1-3)
2001 - Paracorallicarcinus arcanus Tessier, Beschin, Busulini &
De Angeli in De Angeli & Beschin; p. 38, Text-fig. 32
2004 - Paracorallicarcinus arcanus Tessier, Beschin, Busulini &
De Angeli in Beschin & De Angeli; p. 21
Holotype: MCZ 1595 (I.G. 284662).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Tessier et al. (1999) reported 14 specimens
(MCZ from 1584 to 1590, from 1592 to 1598) from
“Main” quarry of Arzignano (Vicenza).
Simonellia Vinassa de Regny, 1 897
Type species: Simonellia quiricensis Vinassa de
Regny, 1897.
Stratigraphic range: Pliocene.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
67
Simonellia quiricensis Vinassa de Regny, 1 897
1897 - Simonellia quiricensis Vinassa de Regny; p. 2, PI. 2 (fig. 1)
1929 - Simonellia quiricensis Vinassa de Regny in Glaessner; p. 378
1969 - Simonellia quiricensis Vinassa de Regny in Glaessner;
p. 521
Holotype: lost.
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Vinassa de Regny (1897) reported one
specimen from S. Quirico d’Orcia (Siena), today lost.
Note: Glaessner (1929) ascribed Simonellia in dubi¬
tative form to thè family Goneplacidae. Later Glaessner
(1969) included this genus in thè family Xanthidae, point-
ing out that it differs from Ch/inocepha/us in fiat smooth
carapace, shape of stemum, and abdomen, and from
Quadrello in shorter anterolateral margin and front.
Family Hexapodidae Miers, 1886
Stevea Manning & Holthuis, 1981
Type species: Hexapus williamsi Glassell, 1938.
Stratigraphic range: middle Eocene (Lutetian) -
Recent.
Stevea cesarli Beschin, Busulini, De Angeli & Tessier,
1994
1994 - Stevea cesarii Beschin, Busulini, De Angeli & Tessier; p. 192,
Text-fig. 8, PI. 10 (figs. 1-5)
2001 - Stevea cesarii Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 38
2004 - Stevea cesarii Beschin, Busulini, De Angeli & Tessier in
Beschin & De Angeli; p. 21
2006 - Stevea cesarii Beschin, Busulini, De Angeli & Tessier in
Guinot; p. 560, 567
Holotype: MCZ 1452 (I.G. 286326).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported six specimens
(MCZ 1176, 1430, from 1451 to 1454) from “Albanello”
and “Boschetto” quarries of Nogarole Vicentino (Vi¬
cenza).
Family Menippidae Ortmann, 1893
Eriphia Latreille, 1817
Type species: Cancer spinifrons Herbst, 1782.
Stratigraphic range: Oligocene - Recent.
Eriphia cocchii Ristori, 1886
1886 - Eriphia cocchii Ristori; p. 105, PI. 2 (figs. 3, 4, 11, 13), PI. 3
(figs. 1-2)
1886 - Eriphia punctulata Ristori; p. 109, PI. 2 (figs. 2, 15-16), nov.
syn.
1 89 1 a - Eriphia punctulata Ristori in Ristori; p. 9
1 9 1 0 - Eriphia cocchii Ristori in Fucini; p. 3, Text-figs. 1 a-c, 2
1929 - Eriphia cocchii Ristori in Glaessner; p. 149
1981 - Eriphia cocchii Ristori in Delle Cave; p. 46
1981 - Eriphia puctulata Ristori in Delle Cave; p. 47
Syntypes: IGF 617E (figured by Ristori in PI. 2, figs.
3, 4, 13, 14); IGF 868E (figured by Ristori in PI. 3, figs.
1-2); IGF 869E and IGF 870E (not figured by Ristori).
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (1886) reported three specimens
from Montrappoli Val d’Era and Montebicchieri (S.
Miniato al Tedesco) (IGF); Ristori (189 la) reported one
specimen from Poggio all’Olio (Empoli) (IGF); Fucini
(1910) reported one specimen from Spicchio (Empoli)
(MSNTC, catalogue number unknown).
Note: Ristori (1886) reported one specimen from
Montrappoli Val d’Era, ascribing it to E. punctulata
(holotype: IGF 614E, figured by Ristori in PI. 2, figs. 2,
14-15), today synonym of E. cocchii.
Eriphia spinifrons (Elerbst, 1782)
1782 - Cancer spinifrons Herbst; p. 185, PI. 1 1 (fig. 65)
1816 - Eriphia spinifrons (Herbst) in Risso; p. 13
1825 - Eriphia spinifrons (Herbst) in Desmarest; p. 126, PI. 14 (fig. 1)
1829 - Eriphia spinifrons (Herbst) in Holl; p. 145
1 865 - Cancer spinifrons Herbst in A. Milne Edwards; p. 305
1885 - Eriphia spinifrons (Herbst) in Carus; p. 514
1911 - Eriphia spinifrons (Herbst) in Magri; p. 13
1914 - Eriphia spinifrons (Herbst) in M. Gemmellaro; p. 87, PI. 1
(fig- 21)
1929 - Eriphia spinifrons (Herbst) in Glaessner; p. 149
1971 - Eriphia spinifrons (Herbst) in Philippe & Secretan; p. 12, PI. B
(figs. 10-11)
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported some
specimens from Monte Pellegrino, Ficarazzi, and Baghe-
ria (Palermo) housed in Marchese di Monterosato ’s col-
lection (repository and catalogue number unknown) and
MGUP (catalogue number unknown).
Note: this species is also reported from thè Miocene
of France.
Eriphia sp.
1846 - Eriphia sp. in E. Sismonda; p. 69, PI. 3 (fig. 6)
1929 -Eriphia sp. in Glaessner; p. 150
Stratigraphic range: middle Miocene (Helvetian).
Occurrence: Piemonte.
Material: E. Sismonda (1846) reported one speci¬
men from Torino (repository and catalogue number
unknown).
Eriphia sp.
1888 - Eriphia sp. in Ristori; p. 215
1929 - Eriphia sp. in Glaessner; p. 150
Stratigraphic range: Miocene.
Occurrence: Toscana.
68
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Material: Ristori (1888) reported this genus from S.
Benedetto in Val Benedetta (Livorno), today lost.
Eriphia sp.
1 889 - Eriphia sp. in Ristori; p. 400
1929 - Eriphia sp. in Glaessner; p. 150
1974 - Eriphia sp. in Mastrorilli; p. 4
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Piemonte.
Material: Ristori (1889) reported two specimens from
Mioglia (Savona), today lost.
Eriphia sp.
1895 - Eriphia sp. in Crema; p. 677
1929 - Eriphia sp. in Glaessner; p. 150
Stratigraphic range: Miocene.
Occurrence: Piemonte.
Material: Crema (1895) reported many specimens
from Torino, today lost.
Eriphia sp.
1975 - Eriphia sp. in Secretan; p. 362, Text-fig. 20, PI. 23 (fig. 1)
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported one specimen
(MSNV 4) from Monte Bolca (Verona).
Eriphia sp.
2004 - Eriphia sp. in Garassino, De Angeli, Gallo & Pasini; p. 271,
Text-fig. 12 a-b
Stratigraphic range: Pliocene.
Occurrence: Piemonte.
Material: Garassino et al (2004) reported one speci¬
men (PU 41180) from Orta San Giulio (Novara).
Family Panopeidae Ortmann, 1893
Bittneria Schweitzer & Karasawa, 2004
Type species: Palaeograpsus attenuatus Bittner, 1875.
Stratigraphic range: middle Eocene (Lutetian).
Bittneria attenuatus (Bittner, 1875)
1875 - Palaeograpsus attenuatus Bittner; p. 100, PI. 2 (fig. 10 a-b)
191 Oa — Palaeograpsus attenuatus Bittner in Fabiani; p. 28
1915 - Palaeograpsus attenuatus Bittner in Fabiani; p. 285
1929 - Palaeograpsus attenuatus Bittner in Glaessner; p. 295
1994 - Palaeograpsus attenuatus Bittner in Beschin, Busulini,
De Angeli & Tessier; p. 195, PI. 9 (fig. 4)
2001 - Palaeograpsus attenuatus Bittner in De Angeli & Beschin;
p. 38
2004 - Bittneria attenuatus (Bittner) in Schweitzer & Karasawa; p. 80,
Text-fig. 1 (6)
2004 - Bittneria attenuatus (Bittner) in Beschin & De Angeli; p. 21
Holotype: MNHB MB. A. 663.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875) reported one specimen
(MNHB MB. A. 663) from S. Giovanni Ilarione (Verona);
Beschin et al. (1994) reported three specimens (MCZ
from 1422 to 1424) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza).
Carinocarcinus Lòrenthey, 1898
Type species: Carinocarcinus zitteli Lòrenthey, 1898.
Stratigraphic range: middle Eocene (Lutetian).
Carinocarcinus zitteli Lòrenthey, 1 898
1898 - Carinocarcinus Zitteli Lòrenthey; p. 138, PI. 10 (fig. 1)
1925 - Carinocarcinus Zitteli Lòrenthey in Schlosser; p. 143
1929 - Carinocarcinus Zitteli Lòrenthey in Glaessner; p. 1 12
1939 - Carinocarcinus Zitteli Lòrenthey in Vecsey; p. 37, Text-
figs. 5-6
1969 - Carinocarcinus zitteli Lòrenthey in Via Boada; p. 369
1969 - Carinocarcinus zitteli Lòrenthey in Glaessner; p. 526, Text-fig.
333 (7)
2005 - Carinocarcinus zitteli Lòrenthey in Beschin, De Angeli, Chec¬
chi & Zarantonello; p. 26, Text-fig. 17, PI. 5 (figs. 1, 5)
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported two speci¬
mens (MV 44, 04/17) from Grola di Comedo Vicentino
(Vicenza).
Note: this species is also reported from middle Eocene
of Germany and Hungary.
Glyphithyreus Reuss, 1859
Type species: Glyphithyreus wetherelli Bell, 1858.
Stratigraphic range: Paleocene - Oligocene.
Glyphithyreus ellipticus (Bittner, 1875)
1875 - Plagiolophus ellipticus Bittner; p. 96, PI. 2 (fig. 8 a-b)
1897 - Plagiolophus ellipticus Bittner in Oppenheim; p. 210
191 Oa — Plagiolophus ellipticus Bittner in Fabiani; p. 26
1915 - Plagiolophus ellipticus Bittner in Fabiani; p. 285
1929 - Plagiolophus ellipticus Bittner in Glaessner; p. 329
2001 - Glyphithyreus ellipticus (Bittner) in De Angeli & Beschin; p.
37
2004 - Glyphithyreus ellipticus (Bittner) in Beschin & De Angeli; p. 21
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875) reported this species from
Brusaferri of Bolca (Verona) (repository and catalogue
number unknown).
Lophopanopeus Rathbun, 1 898
Type species: Xantho bella Stimpson, 1860.
Stratigraphic range: middle Eocene (Lutetian) -
Recent.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
69
Lophopanopeus sp.
1998 - Lophopanopeus sp. in Beschin, Busulini, De Angeli, Tessier &
Ungaro; p. 28, Text-figs. 14-15 (2)
2000 - Lophopanopeus sp. in Beschin, De Angeli & Alberti; p. 15
2001 - Lophopanopeus sp. in De Angeli & Beschin; p. 36
2004 - Lophopanopeus sp. in Beschin & De Angeli; p. 2 1
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1998) reported one speci¬
men (MCZ 1540) from “Rossi” quarry of Monte di Malo
(Vicenza).
Palaeograpsus Bittner, 1875
Type species: Paleograpsus inflatus Bittner, 1875.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Palaeograpsus inflatus Bittner, 1875
1875 - Palaeograpsus inflatus Bittner; p. 100, PI. 2 (fig. 11 a-b)
191 Oa - Palaeograpsus inflatus Bittner in Fabiani; p. 12, 28, PI. 2
(figs. 5-6)
1915 - Palaeograpsus inflatus Bittner in Fabiani; p. 285
1 929 - Palaeograpsus inflatus Bittner in Glaessner; p. 295
1929 - Palaeograpsus inflatus Bittner in Lòrenthey & Beurlen; p. 254,
PI. 16 (fig. 2)
1 994 - Palaeograpsus inflatus Bittner in Beschin, Busulini, De Angeli
& Tessier; p. 194, PI. 9 (fig. 5)
1995 - Palaeograpsus inflatus Bittner in De Angeli; p. 16, Text-fig. 3
(2-3), PI. 2 (figs. 2-4)
1998 - Palaeograpsus inflatus Bittner in Beschin, Busulini, De Angeli,
Tessier & Ungaro; p. 30, Text-fig. 16(1)
2000 - Palaeograpsus inflatus Bittner in Beschin, De Angeli & Alberti;
p. 15
2001 - Palaeograpsus inflatus Bittner in De Angeli & Beschin;
p. 38
2001 - Palaeograpsus inflatus Bittner in Karasawa & Kato; p. 272
2004 - Palaeograpsus inflatus Bittner in Beschin & De Angeli;
p. 21
2004 - Palaeograpsus inflatus Bittner in Schweitzer & Karasawa;
p. 80
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian) -
upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Bittner (1875) reported this species from
Laverda (Vicenza) and Fumane di Polesella (Verona)
without stratigraphic indication (repository and cata-
logue number unknown); Fabiani (191 Oa) reported three
specimens from Bocca d’Ansiesa and “Fontanella” di
Grancona (Vicenza) (repository and catalogue number
unknown); Beschin et al. (1994) reported one specimen
(MCZ 1421) from “Boschetto” quarry of Nogarole Vice¬
ntino (Vicenza); De Angeli (1995) reported 18 specimens
(MCZ from 1498 to 1514) from “Fontanella” di Grancona
(Vicenza); Beschin et al. (1998) reported one specimen
(MCZ 1521) from “Rossi” quarry of Monte di Malo
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Palaeograpsus sp.
1905 - Palaeograpsus sp. in Checchia-Rispoli; p. 322
1929 - Palaeograpsus sp. in Glaessner; p. 296
1933 - Palaeograpsus sp. in Di Salvo; p. 38
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1905) reported one spec¬
imen from Balzo del Gatto (Palermo) (MGUP, catalogue
number unknown); Di Salvo (1933) reported one speci¬
men from La Pietra Lunga (Palermo) (MGUP, catalogue
number unknown).
Panopeus H. Milne Edwards, 1834
Type species: Panopeus herbstii FI. Milne Edwards,
1834.
Stratigraphic range: Eocene - Recent.
Panopeus bolcensis De Zigno, 1915
Cancer bolcensis De Zigno in letteris
1975 - Panopeus bolcensis De Zigno in Secretan; p. 359, Text-fig. 19,
PI. 22, 23 (figs. 3-4)
Holotype: MNHN 18.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported three specimens
(MNHN 18; MGPD 6793-6803, 100-lOObis) from Monte
Bolca (Verona).
Panopeus granulineatus Miiller & Collins, 1991
1933 - Pilodius mediterraneus Lòrenthey in Di Salvo; p. 31, PI. 2
(fig. 6 a-b)
1991 - Panopeus granulineatus Miiller & Collins; p. 74, Text-
fig. 4 d, PI. 5 (figs. 3-4, 6)
Holotype: MAFI EGA- 11.1 (M.91-167).
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported one specimen
(MGUP 2621) from La Pietra Lunga (Palermo).
Note: Di Salvo (1933) included thè Sicilian specimen
in Pilumnus mediterraneus (= Pilodius mediterraneus )
Lòrenthey, 1897, known from thè middle Miocene of
Hungary. The study of thè dorsal surface of thè carapace
of thè Sicilian specimen pointed out shape and characters
similar with Panopeus granulineatus Miiller & Collins,
1991, from thè upper Eocene (Priabonian) of Hungary.
Panopeus vicentinus (Bittner, 1875)
1875 - Panopaeus Vicentinus Bittner; p. 33, PI. 2 (fig. 7)
1910a - Panopaeus vicentinus Bittner in Fabiani; p. 27
1915 - Panopeus vicetinus Bittner in Fabiani; p. 285
1915 - Panopaeus vicentinus Bittner in Dainelli, p. 699
1929 - Panopeus vicentinus Bittner in Glaessner; p. 304
2001 - Panopeus vicentinus Bittner in De Angeli & Beschin; p. 36
2004 - Panopeus vicentinus Bittner in Beschin & De Angeli; p. 21
Holotype: unknown.
70
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto, Friuli-Venezia Giulia.
Material: Bittner (1875) reported this species from
Ciupio (Verona) (repository and catalogue number
unknown); Dainelli (1915) reported one specimen
from Buttrio (Udine) (repository and catalogue number
unknown).
Family Pilumnidae Samouelle, 1819
Eohalimede Blow & Manning, 1996
Type species: Eohalimede talleri Blow & Manning,
1996.
Stratigraphic range: middle Eocene (Lutetian).
Eohalimede granosa Beschin, Busulini, De Angeli &
Tessier, 2002
2002 - Eohalimede granosa Beschin, Busulini, De Angeli & Tessier;
p. 19, Text-fig. 14, PI. 4 (figs. 1-2)
2004 - Eohalimede granosa Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 15
2005 - Eohalimede granosa Beschin, Busulini, De Angeli & Tessier in
Beschin, De Angeli, Checchi & Zarantonello; p. 25, PI. 5 (fig. 6)
Holotype: MCZ 2279 (I.G. 296407).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported two speci-
mens (MCZ 2279, 2280) from “Main” quarry of Arzignano
(Vicenza); Beschin et al. (2005) reported one specimen
(MCZ 2341) from Grola di Comedo Vicentino (Vicenza).
Eopilumnus Beschin, Busulini, De Angeli & Tessier,
2002
Type species: Eopilumnus checchii Beschin, Busulini,
De Angeli & Tessier, 2002.
Stratigraphic range: middle Eocene (Lutetian).
Eopilumnus checchii Beschin, Busulini, De Angeli
& Tessier, 2002
2002 - Eopilumnus checchii Beschin, Busulini, De Angeli & Tessier;
p. 18, Text-fig. 13, PI. 3 (figs. 4-5)
2004 - Eopilumnus checchii Beschin, Busulini, De Angeli & Tessier in
Beschin & De Angeli; p. 20
2004 - Eopiluimnus checchii Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 1 1 5
Holotype: MCZ 2276 (I.G. 296404).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2002) reported three speci-
mens (MCZ from 2276 to 2278) from “Main” quarry of
Arzignano (Vicenza).
Eumorphactaea Bittner, 1875
Type species: Eumorphactaea scissifrons Bittner,
1875.
Stratigraphic range: middle Eocene (Lutetian).
Eumorphactaea scissifrons Bittner, 1875
1875 - Eumorphactaea scissifrons Bittner; p. 92, PI. 2 (fig. 12)
1883 - Eumorphactaea scissifrons Bittner in Bittner; p. 313, PI. 1
(fig. 10)
191 Oa - Eumorphactaea scissifrons Bittner in Fabiani; p. 27
1915 - Eumorphactaea scissifrons Bittner in Fabiani; p. 285
1929 - Eumorphactaea scissifrons Bittner in Glaessner; p. 170
1969 - Eumorphactaea scissifrons Bittner in Glaessner; p. 517
1969 - Eumorphactaea scissifrons Bittner in Via Boada; p. 376
2001 - Eumorphactaea scissifrons Bittner in De Angeli & Beschin;
p. 36
2005 - Eumorphactaea scissifrons Bittner in Beschin, De Angeli,
Checchi & Zarantonello; p. 24, Text-fig. 16, PI. 5 (figs. 2-3)
Holotype: MNHB K114, MB. A. 664.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875, 1883) reported one speci¬
men (MNHB K114, MB.A. 664) from Ciupio (Verona);
Beschin et al. (2005) reported four specimens (MCZ
2354, 2377; MV 45, 05/20) from Grola di Comedo Vi¬
centino (Vicenza).
Galenopsis A. Milne Edwards, 1 865
Type species: Galenopsis typica A. Milne Edwards,
1865.
Stratigraphic range: Eocene - Pliocene.
Galenopsis crassifrons A. Milne Edwards, 1865
1865 - Galenopsis crassifrons A. Milne Edwards; p. 347, PI. 30
(fig- 2)
1901 - Galenopsis crassifrons A. Milne Edwards in Oppenheim;
p. 284
1908 - Galenopsis crassifrons A. Milne Edwards in Fabiani; p. 211,
236
191 Oa — Galenopsis crassifrons A. Milne Edwards in Fabiani; p. 27
191 5 - Galenopsis crassifrons A. Milne Edwards in Fabiani; p. 285
1929 - Galenopsis crassifrons A. Milne Edwards in Glaessner;
p. 177
2001 - Galenopsis crassifrons A. Milne Edwards in De Angeli &
Beschin; p. 37
2004 - Galenopsis crassifrons A. Milne Edwards in Beschin &
De Angeli; p. 2 1
Holotype: MNHN R03798.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: A. Milne Edwards (1862-1865) reported
one specimen (MNHN R03798) from Lonigo (Vicenza).
Galenopsis cfr. G. murchisoni A. Milne Edwards, 1865
1905 - Galenopsis cfr. G. murchisoni A. Milne Edwards in Checchia-
Rispoli; p. 318
1929 - Galenopsis cfr. G. murchisonh A. Milne Edwards in Glaessner;
p. 178
1933 - Galenopsis cfr. G. murchisoni A. Milne Edwards in Di Salvo;
p. 34
Stratigraphic range: middle/upper Eocene.
Occurrence: Sicilia.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DEC APODA FROM ITALY
71
Material: Checchia-Rispoli (1905) reported one spec¬
imen (MGUP 2607) from Balzo del Gatto (Palermo); Di
Salvo (1933) reported one specimen from Balzo del Gatto
(Palermo) (MGUP, catalogue number unknown).
Galenopsis ristori i Checchia-Rispoli, 1907
1907 - Galenopsis ristorii Checchia-Rispoli; p. 26, PI. 1 (figs. 1-2)
1929 - Galenopsis ristorii Checchia-Rispoli in Glaessner; p. 178
1933 - Galenopsis ristorii Checchia-Rispoli in Di Salvo; p. 37
Holotype: MGUP.
Stratigraphic range: middle/upper Eocene.
Occurrence: Sicilia.
Material: Checchia-Rispoli (1907) reported one
specimen from Bagheria (Palermo) (MGUP, catalogue
number unknown); Di Salvo (1933) reported one
specimen from Bichinello (Palermo) (MGUP, catalogue
number unknown).
Galenopsis schopeni Checchia-Rispoli, 1 905
1905 - Galenopsis schopeni Checchia-Rispoli; p. 320, PI. 1
(figs. 3-4)
1929 - Galenopsis schopeni Checchia-Rispoli in Glaessner; p. 179
1933 - Galenopsis schopeni Checchia-Rispoli in Di Salvo; p. 36
Holotype: MGUP.
Stratigraphic range: middle/upper Eocene.
Occurrence: Sicilia.
Material: Checchia-Rispoli (1905) and Di Salvo
(1933) reported four specimens (MGUP 2606, 2608,
2609, 2610) from Balzo del Gatto (Palermo).
Galenopsis similis Bittner, 1875
1875 - Galenopsis similis Bittner; p. 97, PI. 2 (fig. 9)
1898 - Galenopsis similis Bittner in Lòrenthey; p. 64, PI. 5 (figs. 1-2)
1899 - Galenopsis similis Bittner in Oppenheim; p. 58
1905 - Galenopsis similis Bittner in Checchia-Rispoli; p. 319
1910a - Galenopsis similis Bittner in Fabiani; p. 27
1915 - Galenopsis similis Bittner in Fabiani; p. 285
1929 - Galenopsis similis Bittner in Glaessner; p. 179
1929 - Galenopsis similis Bittner in Lòrenthey & Beurlen; p. 247,
PI. 16 (figs. 3-4, 6)
1933 - Galenopsis similis Bittner in Di Salvo; p. 34
1969 - Galenopsis similis Bittner in Via Boada; p. 378
1975 - Galenopsis similis Bittner in Miiller; p. 510, 516
1991 - Galenopsis similis Bittner in Miiller & Collins; p. 86, PI. 8
(figs. 4-5)
2000 - Galenopsis similis Bittner in Beschin, Busulini, De Angeli,
Tessier & Ungaro; p. 8, PI. 1 (fig. 5)
2001 - Galenopsis similis Bittner in De Angeli & Beschin; p. 37
2004 - Galenopsis similis Bittner in Beschin & De Angeli; p. 21
Holotype: unknown.
Stratigraphic range: middle/upper Eocene - lower
Oligocene.
Occurrence: Veneto, Sicilia.
Material: Bittner (1875) reported this species from
Monte di Malo and Muzzolon (Vicenza) (catalogue
number unknown); Checchia-Rispoli (1905) and Di
Salvo (1933) reported two specimens (MGUP 2603,
2605) from Balzo del Gatto and Bichinello (Palermo);
Beschin et al. (2000) reported one specimen (MCZ
1742) from “Gecchelina” quarry of Monte di Malo
(Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Galenopsis sp.
1883 - Galenopsis sp. in Bittner; p. 314
1929 - Galenopsis sp. in Glaessner; p. 179
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1883) reported one specimen from
Monte Magrè di Schio (Vicenza) (repository and cata¬
logue number unknown).
Galenopsis sp.
1886 - Galenopsis sp. in Ristori; p. 103, PI. 2 (fig. 7)
1929 - Galenopsis sp. in Glaessner; p. 179
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (1886) reported one specimen from
Orciano (Pisa), today lost.
Galenopsis sp.
1933 - Galenopsis sp. in Di Salvo; p. 37, PI. 2 (fig. 8)
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported one specimen
(MGUP 2599) from Bichinello (Palermo).
Lobogalenopsis Miiller & Collins, 1991
Type species: Galenopsis quadrilobata Lòrenthey,
1897.
Stratigraphic range: upper Eocene (Priabonian).
Lobogalenopsis quadrilobata (Lòrenthey, 1 897)
1897 - Galenopsis quadrilobata Lòrenthey; p. 156
1898a - Galenopsis quadrilobata Lòrenthey in Lòrenthey; p. 100
1898b - Galenopsis quadrilobata Lòrenthey in Lòrenthey; p. 87,
PI. 5 (fig. 3)
1898c - Galenopsis quadrilobata Lòrenthey in Lòrenthey; p. 60,
PI. 5 (fig. 3)
1929 - Galenopsis quadrilobata Lòrenthey in Glaessner; p. 178
1929 - Galenopsis quadrilobata Lòrenthey in Lòrenthey & Beurlen;
p. 249, PI. 16 (fig. 5)
1933 - Galenopsis quadrilobata Lòrenthey in Di Salvo; p. 35, PI. 3
(fig. 3 a-c)
1969 - Galenopsis quadrilobata Lòrenthey in Via Boada; p. 378
1991 - Lobogalenopsis quadrilobata (Lòrenthey) in Miiller & Collins;
p. 88, Text-fig. 5 f, PI. 8 (figs. 8-10)
Holoty pe: MAFI.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported two specimens
(MGUP 323, 2593) from Balzo del Gatto (Palermo).
72
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Lobonotus A. Milne Edwards, 1 864
Type species: Lobonotus sculptus A. Milne Edwards,
1864.
Stratigraphic range: Eocene - Miocene.
Lobonotus cfr. L. orientalis Collins & Morris, 1978
1983 - Lobonotus cfr. orientalis Collins & Morris in Busulini,
Tessier, Visentin, Beschin, De Angeli & Rossi; p. 65, PI. 3
(fig. 4)
2001 - Lobonotus cfr. orientalis Collins & Morris in De Angeli &
Beschin, p. 36
2004 - Lobonotus cfr. orientalis Collins & Morris in Beschin, Busulini,
De Angeli & Tessier; p. 1 1 5
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1983) reported four speci¬
men (*31, *35, *77; MSNVE 10095) from “Main”
quarry of Arzignano (Vicenza).
Note: Lobonotus orientalis Collins & Morris, 1978,
from thè Eocene of Pakistan was recently ascribed to
thè new genus Pakicarcinus Schweitzer, Feldmann &
Gingerich, 2004. The Italian species must be probably
considered a new species of Lobonotus.
Pilumnus Leach, 1815
Type species: Cancer hirtellus Linnaeus, 1761.
Stratigraphic range: Oligocene - Recent.
Pilumnus villosissimus (Rafinesque-Schmaltz, 1814)
1814 - Cancer villosissimus Rafinesque-Schmaltz; p. 20
1 827 - Pilumnus villosus Risso; p. 1 0
1885 - Pilumnus villosus Risso in Carus; p. 514
1914 - Pilumnus villosus Risso in M. Gemmellaro; p. 86, PI. 1
(figs. 19-20)
1918 - Pilumnus hirtellus var. villosus Risso in Pesta; p. 417, Text-
fig. 137
1929 - Pilumnus hirtellus var. villosa Risso in Glaessner; p. 316
1940 - Pilumnus villosus Risso in Bouvier; p. 256
1956 - Pilumnus villosus Risso in Monod; p. 249
1958 - Pilumnus villosus Risso in Holthuis & Gottlieb; p. 97
1965 - Pilumnus villosus Risso in Forest; p. 378
1969 - Pilumnus villosissimus (Rafinesque-Schmaltz) in Zariquiey
Alvarez; p. 392
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: M. Gemmellaro (1914) reported some
specimen from Ficarazzi (Palermo), housed in Marchese
di Monterosato’s collection (MGUP, catalogue number
unknown).
Pilumnus sp.
1981 - Pilumnus sp. in Varola; p. 29, PI. 7 (fig. 3)
Stratigraphic range: Pliocene.
Occurrence. Puglia.
Material: Varola (1981) reported one specimen from
Leuca (Lecce) (GNSL, catalogue number unknown).
Pilumnus sp.
2004 - Pilumnus sp. in Garassino, De Angeli, Gallo & Pasini; p. 273,
Text-fig. 13
Stratigraphic range: Pliocene.
Occurrence: Piemonte.
Material: Garassino et al. (2004) reported two speci¬
men (PU 41183, 41184) from Cossato (Biella).
Titanocarcinus A. Milne Edwards, 1 863
Type species: Titanocarcinus serratifrons A. Milne
Edwards, 1863.
Stratigraphic range: Upper Cretaceous (Senonian) -
Pliocene.
Titanocarcinus aculeatus Busulini, Tessier & Visentin,
1984
1982 - Titanocarcinus affi raulinianus in Busulini, Tessier & Visentin;
p. 82, Text-fig. 3
1984 - Titanocarcinus aculeatus Busulini, Tessier & Visentin; p. 110,
PI. 1 (figs. 1-3), PI. 2 (figs. 1-2), PI. 3 (figs. 1- 2)
2001 - Titanocarcinus aculeatus Busulini, Tessier & Visentin in
De Angeli & Beschin; p. 35
2002 - Titanocarcinus aculeatus Busulini, Tessier & Visentin in Beschin,
Busulini, De Angeli & Tessier; p. 20, Text-fig. 15, PI. 4 (fig. 3)
2004 - Titanocarcinus aculeatus Busulini, Tessier & Visentin in
Beschin & De Angeli; p. 21
2004 - Titanocarcinus aculeatus Busulini, Tessier & Visentin in
Beschin, Busulini, De Angeli & Tessier; p. 1 1 5
Holotype: MSNVE 10441.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1984) reported 20 speci-
mens (from *60 to *70, *150, *178, from *184 to *187;
MSNVE 10441, 10442, 10443) from “Main” quarry of
Arzignano (Vicenza); Beschin et al. (2002) reported two
specimens (MCZ 1188, 1201) from “Main” quarry of
Arzignano (Vicenza).
Titanocarcinus edwardsi (E. Sismonda, 1 846)
1 846 - Xantho edwardsi E. Sismonda; p. 61, PI. 3 (fig. 5)
1863 - Titanocarcinus edw’ardsi (E. Sismonda) in A. Milne Edwards;
p. 35
1864 - Titanocarcinus edwardsi (E. Sismonda) in A. Milne Edwards;
PI. 10 (fig. 3)
1886 - Titanocarcinus edwardsi (E. Sismonda) in Ristori; p. 99, PI. 2
(fig. 9)
1895 - Titanocarcinus edwardsi (E. Sismonda) in Crema; p. 678, Text-
figs. 17-18
1923 - Titanocarcinus edwardsi (E. Sismonda) in Tettoni; p. 162
1929 - Titanocarcinus edwardsi (E. Sismonda) in Glaessner; p. 384
Holotype: lost.
Stratigraphic range: upper Pliocene (Piacentian).
Occurrence: Piemonte, Toscana.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
73
Material: E. Sismonda (1846) reported two speci-
mens from Torino and Asti, today lost; Ristori (1886)
reported two specimens from Asti and Orciano (Pisa),
today lost; Crema (1895) reported three specimens from
Bra (Cuneo), today lost; fettoni (1923) reported one
specimen from S. Venanzio (Modena) (MGUM, cata-
logue number unknown).
Titanocarcìnus euglyphos Bittner, 1875
1875 - Titanocarcìnus euglyphos Bittner; p. 95, PI. 2 (fig. 6)
1 9 1 Oa - Titanocarcìnus euglyphos Bittner in Fabiani; p. 26
Il 91 5 - Titanocarcìnus euglyphos Bittner in Fabiani; p. 285
1929 - Titanocarcìnus euglyphos Bittner in Glaessner; p. 385
1983 - Titanocarcìnus euglyphos Bittner in Busulini, Tessier, Visentin,
IBeschin, De Angeli & Rossi; p. 66, PI. 3 (fig. 1)
1994 - Titanocarcìnus euglyphos Bittner in Beschin, Busulini,
De Angeli & Tessier; p. 189, PI. 9 (fig. 2)
2001 - Titanocarcìnus euglyphos Bittner in De Angeli & Beschin;
p. 35, Text-fig. 31
2004 - Titanocarcìnus euglyphos Bittner in Beschin & De Angeli;
P- 21
2004 - Titanocarcìnus euglyphos Bittner in Beschin, Busulini,
De Angeli & Tessier; p. 1 15
Holotype: unknown.
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875) reported this species from
Ciupio (Verona) (repository and catalogue number
unknown); Busulini et al. (1983) reported seven speci¬
mens (*6, *9, *21, *22, *76; MSN VE 10097, 10098)
from “Main” quarry of Arzignano (Vicenza); Beschin et
al. (1994) reported three specimens (MCZ 1223, 1306,
1457) from “Boschetto” quarry of Nogarole Vicentino
(Vicenza).
Note: Feldmann et al. (1998) described this species
from thè middle Eocene (Lutetian) of N Carolina (United
States). The American specimen show thè following dif-
ferences respect thè Italian ones: wider carapace, more
coverging posterolateral margins and wider cardiac
region. Therefore, thè American species could be ascribed
to a different species.
Titanocarcìnus kochii Lòrenthey, 1 897
1897 - Titanocarcìnus kochii Lòrenthey; p. 155
1929 - Titanocarcìnus kochii Lòrenthey in Lòrenthey & Beurlen;
p. 239, PI. 11 (figs. 4-5)
1929 - Titanocarcìnus kochi Lòrenthey in Glaessner; p. 385
1933 - Titanocarcìnus kochii Lòrenthey in Di Salvo; p. 28, PI. 1
(fig. 2 a-b)
1 969 - Titanocarcìnus kochii Lòrenthey in Via Boada; p. 420
1991 - Titanocarcìnus kochii Lòrenthey in Muller & Collins; p. 81, PI.
6 (figs 2, 5-6)
Holotype: MAFI.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported two specimens
(MGUP 26 1 3, 26 1 8) from La Pietra Lunga and San Cipir-
rello (Palermo).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Titanocarcìnus raulinianus A. Milne Edwards, 1863
1863 - Titanocarcìnus raulinianus A. Milne Edwards; p. 37
1864 - Titanocarcìnus raulinianus A. Milne Edwards in A. Milne
Edwards; PI. 9 (figs. 3-4)
1893 - Titanocarcìnus raulinianus A. Milne Edwards in Ristori; p. 212
1898 - Titanocarcìnus raulinianus A. Milne Edwards in Lòrenthey;
p. 57
1903 - Titanocarcìnus raulinianus A. Milne Edwards in Lòrenthey;
p. 113, PI. 2 (fig. 1)
1929 - Titanocarcìnus raulinianus A. Milne Edwards in Glaessner;
p. 385
2000 - Titanocarcìnus raulinianus A. Milne Edwards in Beschin,
Busulini, De Angeli, Tessier & Ungaro; p. 8, PI. 2 (fig. 3)
2001 - Titanocarcìnus raulinianus A. Milne Edwards in De Angeli &
Beschin; p. 36
2004 - Titanocarcìnus raulinianus A. Milne Edwards in Beschin &
De Angeli; p. 21
2006a - Titanocarcìnus raulinianus A. Milne Edwards in De Angeli &
Garassino; p. 287, Text-fig. 1 1
Holotype: unknown.
Stratigraphic range: Eocene.
Occurrence: Veneto, Friuli-Venezia Giulia, Puglia.
Material: Ristori (1893) reported one specimen
from Monte Saraceno (Gargano) (repository and cata¬
logue number unknown); Beschin et al. (2000) reported
one specimen (MCZ 1819) from “Gecchelina” quarry
of Monte di Malo (Vicenza); De Angeli & Garassino
(2006a) reported one specimen (MFSN 29051) from
Casali Ottelio (Udine).
Note: this species is also reported from thè middle/
upper Eocene of France and Hungary.
Titanocarcìnus subovalis Ristori, 1 896
1896 - Titanocarcìnus subovalis Ristori; p. 504, PI. 12 (figs. 3-4)
1929 - Titanocarcìnus subovalis Ristori in Glaessner; p. 386
1981 - Titanocarcìnus subovalis Ristori in Delle Cave; p. 47
Holotype: IGF 620E (figured by Ristori in PI. 2,
figs. 3-4).
Stratigraphic range: Pliocene.
Occurrence: Toscana.
Material: Ristori (1896) reported one specimen (IGF
620E) from Arbia (Siena).
Titanocarcìnus sp.
1933 - Titanocarcìnus sp. in Di Salvo; p. 30, PI. 2 (fig. 9)
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported two specimens
(MGUP 2617, 2619) from Ponte di Castronuovo and
Balzo del Gatto (Palermo).
Family Xanthidae MacLeay, 1838
Monodaeus Guinot, 1 967
Type species: Xantho couchi Bell in Couch, 1851
(= Xantho tuberculata Bell, 1852).
74
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Stratigraphic range: Pliocene - Recent.
Monodaeus bortolottii Delle Cave, 1988
1988b - Monodaeus bortolottii Delle Cave; p. 123, PI. 1 (figs. 1-2),
PI. 2 (figs. 1-5)
1993 - Monodaeus cfr. M. bortolottii Delle Cave in Miiller; p. 21,
Text-fig. 1 0 F
Holotype: IGF 1786E.
Stratigraphic range: upper Pliocene (Piacentian).
Occurrence: Toscana.
Material: Delle Cave (1988b) reported one specimen
(IGF 1786E) from Botro delTAlpino (Volterra).
Note: this species is also reported from thè Neogene
of Spain.
Paraxanthosia Miiller & Collins, 1 99 1
Type species: Paraxanthosia budensis Miiller & Col¬
lins, 1991.
Stratigraphic range: middle/upper Eocene.
Paraxanthosia tubercu/ata Beschin, De Angeli, Checchi
& Zarantonello, 2005
2005 - Paraxanthosia tuberculata Beschin, De Angeli, Checchi &
Zarantonello; p. 23, Text-fig. 15, PI. 4 (fig. 8)
Holotype: MCZ 2333 (I.G. 296552).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported one speci¬
men (MCZ 2333) from Grola di Comedo Vicentino
(Vicenza).
Phlyctenodes A. Milne Edwards, 1 862
Type species: Phlyctenodes tuberculosus A. Milne
Edwards, 1862.
Stratigraphic range: Eocene - ?Miocene.
Phlyctenodes dalpiazi Fabiani, 1911
191 Oa - Phlyctenodes aff. Phl. Krenneri in Fabiani; p. 25
1911 - Phlyctenodes Dalpiazi Fabiani; p. 42, Text-fig. 1 a-b
1915 - Phlyctenodes Dalpiazi Fabiani in Fabiani; p. 285
1929 - Phlyctenodes dalpiazi Fabiani in Glaessner; p. 3 13
2001 - Phlyctenodes dalpiazi Fabiani in Beschin, De Angeli & Chec¬
chi; p. 29
2001 - Phlyctenodes dalpiazi Fabiani in De Angeli & Beschin;
p. 34
2002 - Phlyctenodes dalpiazi Fabiani in De Angeli & Garassino; p. 5
Holotype: MGPD 23654.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Veneto.
Material: Fabiani (191 Oa, 1911) reported one speci¬
men (MGPD 23654) from Tongriani limestones (Oli¬
gocene) of S. Feliciano (Vicenza).
Note: a recent study on thè limestones including
a decapod fauna of S. Feliciano allowed to date these
levels to thè upper Eocene (Priabonian) (De Angeli &
Garassino, 2002).
? Phlyctenodes hantkeni Lòrenthey, 1 898
1898 - Phlyctenodes Hantkeni Lòrenthey; p. 44, PI. 2 (fig. 10)
1929 - Phlyctenodes Hantkeni Lòrenthey in Lòrenthey & Beurlen;
p. 199, PI. 12 (fig. 8)
1929 - Phlyctenodes Hantkeni Lòrenthey in Glaessner; p. 313
1933 - Phlyctenodes hantkeni Lòrenthey in Di Salvo; p. 21, PI. 2
(fig. 1 a-d)
1991 - 1 Phlyctenodes hantkeni Lòrenthey in Miiller & Collins; p. 49
Holotype: MAFI.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported 13 specimens
(MGUP 321, 423, 2587, 2588, 2589, 2590, 2591, 2595,
2600, 2601) from Balzo del Gatto, La Pietra Lunga, San
Cipriello (Palermo).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
IPhlyctenodes irregu/aris Ristori, 1 896
1896 - Phlyctenodes irregularis Ristori; p. 506, PI. 12 (fig. 1)
1929 - Phlyctenodes irregularis Ristori in Glaessner; p. 313
Holotype: unknown.
Stratigraphic range: middle Miocene.
Occurrence: Piemonte.
Material: Ristori (1896) reported one specimen
from Serravalle (repository and catalogue number
unknown).
Note: Ristori (1896) described this species on thè
morphological characters of one incomplete cara¬
pace poorly preserved, ascribing it to Phlyctenodes
A. Milne Edwards, 1862, for thè presence of strong
irregular tubercles on thè dorsal surface of carapace.
However Phlyctenodes is a typically genus from thè
Eocene and thè ornamentation with strong irregular
tubercles of Ristori’s species resembles that of Daira
speciosa (Reuss, 1871) discovered invariably in reefal
structures in thè Paratetheys and in thè Mediterranean
Miocene.
Phlyctenodes krenneri Lòrenthey, 1 897
1897 - Phlyctenodes Krenneri Lòrenthey; p. 154
1898 - Phlyctenodes Krenneri Lòrenthey; p. 46, PI. 2 (fig. 10)
1905 - Phlyctenodes Krenneri Lòrenthey in Checchia-Rispoli; p. 312,
PI. 1 (fig. 10)
191 1 - Phlyctenodes Krenneri Lòrenthey in Fabiani; p. 44, Text-fig. 2
1929 - Phlyctenodes Krenneri Lòrenthey in Glaessner; p. 3 13
1933 - Phlyctenodes krenneri Lòrenthey in Di Salvo; p. 20
1969 - Phlyctenodes krenneri Lòrenthey in Via Boada; p. 408
1975 - Phlyctenodes krenneri Lòrenthey in Miiller; p. 516, 520
1991 - Phlyctenodes krenneri Lòrenthey in Miiller & Collins; p. 76,
PI. 5 (fig. 9), PI. 6 (fig. 1)
Holotype: MAFI.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1905) reported one
specimen from Balzo del Gatto (Palermo) (MGUP,
catalogue number unknown); Di Salvo (1933) reported
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM 1TALY
75
six specimens (MGUP 2602, 2623, 2596, 2597, 2598)
from Bichinello, La Pietra Lunga, and Balzo del Gatto
(Palermo).
Note: this species is also reported fron thè upper
Eocene (Priabonian) of Hungary.
Phlyctenodes nicolisi Bittner, 1884
1884 - Phlyctenodes Nicolìsi Bittner; p. 17, PI. 1 (fig. 5)
1915 - Phlyctenodes Nicolisi Bittner in Fabiani; p. 284
1 929 - Phlyctenodes Nicolisi Bittner in Glaessner; p. 3 1 3
2000 - Phlyctenodes nicolisi Bittner in Beschin, Busulini, De Angeli,
ITessier & Ungaro; p. 8, PI. 2 (fig. 2 a-b)
2001 - Phlyctenodes nicolisi Bittner in De Angeli & Beschin, p. 34
Holotype: MSNV 1892.
Stratigraphic range: lower/ middle Eocene (Ypresian
- Lutetian).
Occurrence: Veneto.
Material: Bittner (1884) reported one specimen
(MSNV 1892) from “Scole” quarry of Avesa (Verona);
Beschin et al. (2000) reported one specimen (MCZ
1830) from “Gecchelina” quarry of Monte di Malo
(Vicenza).
Phlyctenodes steinmanni Lòrenthey, 1 90 1
1901 - Phlyctenodes Steinmanni Lòrenthey; p. 815, PI. 1 (fig. 4)
1901 - Phlyctenodes Steinmanni Lòrenthey; p. 1 1 1, PI. 1 (fig. 4)
1 1929 - Phlyctenodes Steinmanni Lòrenthey in Glaessner; p. 314
1929 - Phlyctenodes Steinmanni Lòrenthey in Lòrenthey & Beurlen;
J p. 200, PI. 12 (fig. 2)
Il 991 - Phlyctenodes steinmanni Lòrenthey in Miiller & Collins; p. 76,
PI. 5 (fig. 13)
1994 - Phlyctenodes steinmanni Lòrenthey in Beschin, Busulini,
De Angeli & Tessier; p. 188, PI. 9 (fig. 3)
2000 - Phlyctenodes cf. steinmanni Lòrenthey in Beschin, Busulini,
De Angeli, Tessier & Ungaro; p. 8, PI. 2 (fig. 4)
2001 - Phlyctenodes steinmanni Lòrenthey in De Angeli & Beschin;
p. 35
Holotype: MAFI.
Stratigraphic range: middle/upper Eocene (Lutetian -
Priabonian).
Occurrence: Veneto.
Material: Beschin et al. (1994) reported two speci¬
mens (MCZ 1214, 1215) from “Boschetto” quarry of
Nogarole Vicentino (Vicenza); Beschin et al. (2000)
reported one specimen (MCZ 1784) from “Gecchelina”
quarry of Monte di Malo (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Phlyctenodes sp.
1982 - Phlyctenodes sp. in Busulini, Tessier & Visentin; p. 82
2004 - Phlyctenodes sp. in Beschin, Busulini, De Angeli & Tessier;
p. 115
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Busulini et al. (1982) reported one speci¬
men from “Main” quarry of Arzignano (Vicenza) (SV,
catalogue number unknown).
Xantho Leach, 1 804
Type species: Xantho floridus Montagu, 1813.
Stratigraphic range: Miocene - Recent.
Xantho floridus Montagu, 1813
1813 -Xantho floridus Montagu; p. 85, PI. 2 (fig. 1)
1815 - Cancer incisus Leach; p. 320, PI. 1 1 (figs. 1-6)
1815 - Xantho floridus Montagu in Leach; p. 320
1 830 - Cancer floridus (Montagu) in Desmarest & Bosc; p. 205
1 834 - Xantho floridus Montagu in A. Milne Edwards; p. 324
1850 - Cancer poressa Olivi in Costa; p. 8, PI. 1 (fig. 2)
1885 - Xantho floridus Montagu in Carus; p. 512, 513
1899 - Xantho floridus Montagu in Acloque; p. 142
1903 - Xantho floridus Montagu in Checchia-Rispoli; p. 488, Text-fig.
without number
1914 -Xantho floridus Montagu in M. Gemmellaro; p. 58, PI. 1 (fig. 18)
1921 -Xantho floridus Montagu in Bell; p. 9
1929 - Xantho floridus Montagu in Glaessner; p. 394
Holotype: unknown.
Stratigraphic range: upper Pleistocene (Sicilian).
Occurrence: Sicilia.
Material: Checchia-Rispoli (1903) reported one
specimen from Palermo, housed in Di-Stefano’s col-
lection (repository and catalogue number unknown);
M. Gemmellaro (1914) reported some specimens
from Ficarazzi (Palermo), housed in Marchese di
Monterosato’s collection (MGUP, catalogue number
unknown).
IXantho lovisatoi (Lòrenthey, 1907)
1907 -Xanthus lovisatoi Lòrenthey; p. 91, PI. 3 (fig. 12)
1909 - IXanthus lovisatoi Lòrenthey in Lòrenthey; p. 245, PI. 1
(fig. 12 a-d)
1929 - IXantho lovisatoi (Lòrenthey) in Glaessner; p. 395
Holotype: unknown.
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Lòrenthey (1909) reported some speci¬
mens from Capo S. Elia (Cagliari), housed in Lovi-
sato’s collection (repository and catalogue number
unknown).
Xantho moldavicus (Yanakevich, 1977)
1928 - Titanocarcinus vulgaris Glaessner; p. 185, PI. 3 (figs. 9-11)
1929 - Titanocarcinus vulgaris Glaessner in Glaessner; p. 386
1953b - Titanocarcinus vulgaris Glaessner in Bachmayer; p. 254, PI. 4
(figs. 1-9), PI. 6 (figs. 1-2)
1974 - Xantho cfr. X. incisus Leach in Miiller; p. 123, PI. 3 (figs. 1-2)
1976 - Xantho incisus Leach in Miiller; p. 152
1977 - Medaeus moldavicus Yanakevich; p. 80, PI. 10 (fig. 4)
1979 - Xantho cfr. X. incisus Leach in Miiller; p. 274, PI. 20 (figs. 1-5)
1979 - Xantho cfr. X. vulgaris (Glaessner) in Forster; p. 100, Text-
figs. 14-15, PI. 3 (figs. 1-3), PI. 4 (figs. 1-4)
1984 -Xantho moldavicus (Yanakevich) in Miiller; p. 92, Text-figs. 5-8,
PI. 86 (figs. 1-5), PI. 87 (fig. 1)
1987 - Xantho cf. moldavicus (Yanakevich) in Friebe; p. 61, PI. 2
(fig- 5)
1991 -Xantho moldavicus (Yanakevich) in Marras & Ventura; p. 1 10
76
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1996 -Xantho moldavicus (Yanakevich) in Miiller; p. 1 1, PI. 2 (fig. 6)
1998 - Xantho moldavicus (Yanakevich) in Muller; p. 34
2006 - Xantho moldavicus (Yanakevich) in Radwanski, Górka &
Wysocka; p. 96, PI. 2 (figs. 5, 6)
Holotype: MHMW 1927/1/3.
Stratigraphic range: Miocene.
Occurrence: Sardegna.
Material: Marras & Ventura (1991) reported one
specimen from Sassari (reposiiory and catalogue number
unknown).
Note: this species is also reported from thè lower to
thè upper Miocene of Austria, Hungary, Poland, Molda¬
via, and Spain.
Xantho sp.
2001 - Xantho sp. in Beschin, De Angeli & Checchi; p. 26, PI. 3
(fig- 4)
2001 -Xantho sp. in De Angeli & Beschin; p. 35
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported one specimen
(MCZ 2119) from Castelgomberto (Vicenza).
f Family Zanthopsidae Via Boada, 1959
Harpactocarcinus A. Milne Edwards, 1 862
Type species: Cancer punctulatus Desmarest, 1822.
Stratigraphic range: Eocene.
Harpactocarcinus macrodactylus (H. Milne Edwards,
1850)
1850 - Cancer macrodactylus H. Milne Edwards in D’Archiac;
p. 304
1862 - Harpactocarcinus macrodactylus (H. Milne Edwards) in A.
Milne Edwards; p. 70, PI. 10 (figs. 1, 1 a)
1875 - Harpactocarcinus macrodactylus (H. Milne Edwards) in Bitt-
ner; p. 87
1929 - Harpactocarcinus punctulatus (Desmarest) in Glaessner;
p. 204
1969 - Harpactocarcinus punctulatus (Desmarest) in Via Boada;
p. 381
2000 - Harpactocarcinus macrodactylus (H. Milne Edwards) in
Beschin, De Angeli & Alberti; p. 1 5
2001 - Harpactocarcinus macrodactylus (H. Milne Edwards) in
De Angeli & Beschin, p. 33, Text-fig. 29
2003 - Harpactocarcinus punctulatus (Desmarest) in Schweitzer;
p. 1118
Holotype: MNHN R03825.
Stratigraphic range: lower Eocene (Ypresian).
Occurrence: Veneto.
Material: H. Milne Edwards (1850) and A. Milne
Edwards (1862-1865) reported one specimen (MNHN
R03825) from San Floriano (Verona).
Note: this species was described by H. Milne Edwards
(1850), and figured later by A. Milne Edwards (1862) on
only one male specimen with a cheliped strongly devel-
oped from thè lower Eocene of S. Floriano (Verona).
Glaessner (1929, p. 204), Via Boada (1969, p. 253),
and Schweitzer (2003, p. 1118) suggested this species
as synonym with H. punctulatus (Desmarest, 1822).
However, H. macrodactylus differs from H. punctulatus
because thè narrower frontal lobes and thè different ratio
between thè fronto-orbital margin and thè maximum wide
of thè carapace. The strong development of thè cheliped
and thè evident convexity of thè lower margin of thè
fixed finger, considered as probable anomaly of thè type
specimen, is instead a typical morphological character of
this species. Harpactocarcinus macrodactylus , from thè
lower/middle Eocene of Monte Baldo and Lessini Vero¬
nesi, is sometimes housed in some Museums of Veneto
with thè name H. punctulatus. Three specimens (MCZ
1319, 1320, 1357) of this species come from Ferrara del
Baldo (Verona).
Harpactocarcinus ova/is A. Milne Edwards, 1 862
1862 - Harpactocarcinus ovalis A. Milne Edwards; p.72, PI. 9
(fig- 2)
1 875 - Harpactocarcinus ovalis A. Milne Edwards in Bittner; p. 88
191 Oa — Harpactocarcinus ovalis A. Milne Edwards in Fabiani; p. 26
1915 - Harpactocarcinus ovalis A. Milne Edwards in Fabiani;
p. 284
1927 - Harpactocarcinus ovalis A. Milne Edwards in Van Straelen;
p. 89
1929 - Harpactocarcinus ovalis A. Milne Edwards in Glaessner;
p. 203
2003 - Harpactocarcinus ovalis A. Milne Edwards in Schweitzer;
p. 1118
Holotype: MNHN R03816 (syntype, cast).
Stratigraphic range: Eocene.
Occurrence: Veneto.
Material: Bittner (1875) reported this species from
Vicentino (locality, repository and catalogue number
unknown).
Note: this species is also reported from thè Eocene of
Spain.
Harpactocarcinus punctulatus (Desmarest, 1822)
1 820 - Brachyurites australis in Schlotheim; p. 36 (nom. nud.)
1822 - Cancer punctulatus Desmarest; p. 92, PI. 7 (figs. 3-4)
1846 - Cancer punctulatus Desmarest in d’Archiac; p. 216
1 850 - Cancer punctulatus Desmarest in H. Milne Edwards; p. 304
1 852 - Cancer punctulatus Desmarest in v. Meyer; p. 302
1 854 - Cancer punctulatus Desmarest in Catullo; p. 886
1 859 - Cancer punctulatus Desmarest in Reuss; p. 25, PI. 1 5 (figs. 1 -5),
PI. 16 (figs. 1-4), PI. 17 (figs. 1-4)
1859 - Cancer brachychelus Reuss; p. 29, PI. 13 (fig. 5), PI. 18
(figs. 1-3)
1861 - Cancer punctulatus Desmarest in Michelotti; p. 139
1862 - Harpactocarcinus punctulatus (Desmarest) in A. Milne
Edwards; p. 66, PI. 8 (fig. 1), PI. 9 (fig. 1)
1875 - Harpactocarcinus punctulatus (Desmarest) in Bittner; p. 86
1 883 - Harpactocarcinus punctulatus (Desmarest) in Bittner; p. 3 1 1
1895 - Cancer ( Harpactocarcinus ) punctulatus (Desmarest) in
De Gregorio; p. 10, PI. 1 (figs. 1-4), PI. 2 (figs. 1-2, 4-6)
1898 - Harpactocarcinus punctulatus (Desmarest) in Lòrenthey; p. 13,
78, PI. 7 (fig. 1)
1901 - Harpactocarcinus punctulatus (Desmarest) in Oppenheim;
p. 280
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DEC APODA FROM ITALY
77
1904 - Harpactocarcinus punctulatus (Desmarest) in Checchia-Ris-
poli; p. 50, Text-figs. 1-3
1908 - Harpactocarcinus punctulatus (Desmarest) in Fabiani; p. 210,
236
1909 - Harpactocarcinus punctulatus (Desmarest) in Toniolo; p. 292,
PI. 26 (fig. 3)
191 Oa - Harpactocarcinus punctulatus (Desmarest) in Fabiani; p. 25
1915 - Harpactocarcinus punctulatus (Desmarest) in Fabiani; p. 284,
285
1926 - Harpactocarcinus punctulatus (Desmarest) in Lòrenthey;
p. 19
1927 - Harpactocarcinus punctulatus (Desmarest) in Van Straelen;
p. 88
1929 - Harpactocarcinus punctulatus (Desmarest) in Glaessner;
p. 204
1929 - Harpactocarcinus punctulatus (Desmarest) in Lòrenthey &
Beurlen; p. 206, PI. 11 (fig. 1)
1936 - Harpactocarcinus punctulatus (Desmarest) in Van Straelen;
p. 5
1941 - Harpactocarcinus punctulatus (Desmarest) in Via Boada; p. 39,
Pls. 4, 8,9, 11
1952 - Harpactocarcinus punctulatus (Desmarest) in Via Boada; p. 80
1953 - Harpactocarcinus punctulatus (Desmarest) in Roger; p. 356,
PI. 10 (fig. 12)
1959 - Harpactocarcinus punctulatus (Desmarest) in Via Boada;
p. 381
1962 - Harpactocarcinus punctulatus (Desmarest) in Piccoli & Mocel-
lin; p. 65
1969 - Harpactocarcinus punctulatus (Desmarest) in Via Boada;
p. 239, Text-fig. 29, Pls. 24-27
2000 - Harpactocarcinus punctulatus (Desmarest) in Beschin,
De Angeli & Alberti; p. 15
2001 - Harpactocarcinus punctulatus (Desmarest) in De Angeli &
Beschin; p. 33
2003 - Harpactocarcinus punctulatus (Desmarest) in Schweitzer;
p. 1118
2004 - Harpactocarcinus punctulatus (Desmarest) in Beschin &
De Angeli; p. 20
Holotype: unknown.
Stratigraphic range: lower/upper Eocene (Ypresian -
Priabonian).
Occurrence: Veneto.
Material: Desmarest (1822), Catullo (1854), Reuss
(1859), A. Milne Edwards (1862-1865), and Bittner
(1875) reported this species from thè Eocene of some
localities of Veneto (repository and catalogue number
unknown); De Gregorio ( 1 895) reported seven specimens
from Valrovina (Vicenza), Verona area, and Castelrotto di
Valpolicella (repository and catalogue number unknown);
Checchia-Rispoli (1904) reported some specimens from
Peschici (Puglia) (repository and catalogue number
unknown); Fabiani (191 Oa) reported this species from
some localities (Nanto, Mossano, and Barbarano - Monti
Berici; S. Floriano, Lavacille, and Valrovina - Vicenza)
(repository and catalogue number unknown); Piccoli &
Mocellin (1962) reported one specimen (MGPD 1130)
from Priabona (Vicenza).
Note: this species is also reported from thè Eocene of
Spain, France, Hungary, Switzerland, Istria, and Albany.
Harpactocarcinus spec.?
1875 - Harpactocarcinus spec.? in Bittner; p. 88, PI. 3 (fig. 3)
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Bittner (1875) reported one specimen from
Sangonini di Lugo (Vicenza) (repository and catalogue
number unknown).
Harpactoxanthopsis Via Boada, 1959
Type species: Cancer quadrilobatus Desmarest,
1822.
Stratigraphic range: Eocene.
Harpactoxanthopsis quadrilobata (Desmarest, 1 822)
1822 - Cancer quadrilobatus Desmarest; p. 93, PI. 8 (figs. 1-2)
1 846 - Cancer kressenbergensis v. Meyer; p. 462, nov. syn.
1847 - Cancer quadrilobatus Desmarest in Delbos; p. 716
1850 - Cancer quadrilobatus Desmarest in d’Archiac; p. 448
1852 - Cancer kressenbergensis v. Meyer in v. Meyer; p. 32
1959 - Cancer quadrilobatus Desmarest in Reuss; p. 81
1862 - Harpactocarcinus quadrilobatus (Desmarest) in A. Milne
Edwards; p. 74, PI. 3 (fig. 2), PI. 4, (fig. 1), PI. 5 (fig. 1)
1862 - Xanthopsis kressenbergensis (v. Meyer) in v. Meyer; p. 156,
PI. 16 (Figs. 12, 14)
1863 - Xanthopsis kressenbergensis (v. Meyer) in A. Milne Edwards;
p. 323, PI. 5 (fig. 3)
1863 - Xanthopsis kressenbergensis (v. Meyer) in Schafhault; p. 226,
PI. 60 (fig. 7)
1864 - Cycloxanthus quadrilobatus (Desmarest) in Jacquot; p. 27
1875 - Harpactocarcinus quadrilobatus (Desmarest) in Bittner; p. 89,
PI. 2 (figs. 4-5), PI. 3 (figs. 1-2)
1883 - Harpactocarcinus quadrilobatus (Desmarest) in Bittner; p. 312
1895 - Cancer (Palaeocarpilius) gecchelinensis De Gregorio; p. 14,
PI. 4 (fig. 3), nov. syn.
1898 - Harpactocarcinus quadrilobatus (Desmarest) in Lòrenthey;
p. 12
1905 - Xanthopsis kressenbergensis (v. Meyer) in Airaghi; p. 206,
PI. 4 (figs. 4-5)
1909 - Harpactocarcinus quadrilobatus (Desmarest) in Toniolo;
p. 292, PI. 26 (fig. 4)
191 Oa - Harpactocarcinus quadrilobatus (Desmarest) in Fabiani;
p. 25
191 Oa - Xanthopsis kressenbergensis (v. Meyer) in Fabiani; p. 26
1912 - Harpactocarcinus quadrilobatus (Desmarest) in Vogl; p. 106
1915 - Harpactocarcinus quadrilobatus (Desmarest) in Fabiani;
p. 284, 285
1915 - Xanthopsis kressenbergensis (v. Meyer) in Fabiani; p. 285
1925 - Xanthopsis kressenbergensis (v. Meyer) in Schlosser; p. 144
1929 - Harpactocarcinus quadrilobatus (Desmarest) in Glaessner;
p. 205
1929 - Xanthopsis quadrilobata (Desmarest) in Lòrenthey & Beurlen;
p. 208, PI. 9 (figs. 3-4), PI. 10 (fig. 7)
1959 - Harpactoxanthopsis quadrilobata (Desmarest) in Via Boada;
p. 54
1962 - Xanthopsis kressenbergensis (v. Meyer) in Piccoli & Mocellin;
p. 77, PI. 4 (fig. 18)
1968 - Harpactoxanthopsis quadrilobatus (Desmarest) in Vogeltanz;
P- 37
1969 - Harpactoxanthopsis quadrilobata (Desmarest) in Via Boada;
p. 276, PI. 30 (figs. 1-2), PI. 31 (figs. 1-2) PI. 32 (fig. 33)
1 969 - Harpactoxanthopsis kressenbergensis (v. Meyer) in Via Boada;
p. 284
1982 - Harpactoxanthopsis quadrilobata (Desmarest) in Busulini,
Tessier & Visentin; p. 80
78
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
1989 - Harpactoxanthopsis quadrilobata (Desmarest) in Solè & Via
Boada; p. 29
1994 - Harpactoxanthopsis quadrilobata (Desmarest) in Beschin,
Busulini, De Angeli & Tessier; p. 1 86, PI. 8 (fig. 1 a-b)
1998 - Harpactoxanthopsis quadrilobata (Desmarest) in Beschin,
Busulini, De Angeli, Tessier & Ungaro; p. 24, Text-figs. 9 (5), 12,
13, 15(1)
1998 - Harpactoxanthopsis quadrilobata (Desmarest) in Miiller;
p. 33
2000 - Harpactoxanthopsis quadrilobata (Desmarest) in Beschin,
De Angeli & Alberti; p. 15
2000 - Harpactoxanthopsis kressenbergensis (v. Meyer) in Beschin,
De Angeli & Alberti; p. 15
2001 - Harpactoxanthopsis quadrilobata (Desmarest) in De Angeli &
Beschin; p. 34, Text-figs. 28-29
2001 - Harpactoxanthopsis kressenbergensis (v. Meyer) in De Angeli
& Beschin; p. 34
2003 - Harpactoxanthopsis quadrilobatus (Desmarest) in Schweitzer;
p. 1119
2004 - Harpactoxanthopsis quadrilobata (Desmarest) in Beschin,
Busulini, De Angeli & Tessier; p. 1 15
2004 - Harpactoxanthopsis quadrilobata (Desmarest) in Beschin &
De Angeli; p. 20
2005 - Harpactoxanthopsis quadrilobata (Desmarest) in Beschin,
De Angeli, Checchi & Zarantonello; p. 23
Holotype: MNHN R03824 (syntype), MNHN R03817
(figured).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material; Desmarest (1822), Reuss (1859), A.
Milne Edwards (1862-1865), Bittner (1875, speci¬
men from Bittner, 1875 = MNHB MB. A. 662), and
De Gregorio (1895) reported this species from thè
Eocene of some localities of Veneto (repository and
catalogue number unknown); Airaghi (1905) reported
two specimens ( H . kressembergensis ) (MGPD, 7476)
from Priabona and Val di Lonte (=Val di Lonta in Ai¬
raghi) (Vicenza); Piccoli & Mocellin (1962) reported
one specimen (MGPD 1123) from Valico di Priabona
(Vicenza); Busulini et al. (1982) reported two speci¬
mens from “Main” quarry of Arzignano (Vicenza)
(repository and catalogue number unknown); Beschin
et al. (1994) reported three specimens (MCZ 1200,
1432, 1450) from “Boschetto” quarry of Nogarole
Vicentino (Vicenza); Beschin et al. (1998) reported
nine specimens (MCZ 1198, 1199, 1257, 1355, 1356,
1528, 1529, 1530, 1531) from “Rossi” quarry of Monte
di Malo (Vicenza).
Note: this species is also reported from thè Eocene of
Spain, France, Germany, Austria, Hungary, Croatia, and
Albany.
Harpactoxanthopsis cfr. H. quadrilobata (Desmarest,
1822)
2005 - Harpactoxanthopsis cfr. H. quadrilobata in Beschin, De Angeli,
Checchi & Zarantonello; p. 23, PI. 4 (fig. 4)
Stratigraphic range; middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (2005) reported two speci¬
mens (MCZ 2311, 2378) from Grola di Comedo Vicen¬
tino (Vicenza).
Harpactoxanthopsis souverbiei (A. Milne Edwards,
1862)
1862 - Harpactocarcinus souverbiei X. Milne Edwards; p. 72, PI. 6
(figs. 3-5)
1875 - Harpactocarcinus souverbiei A. Milne Edwards in Bittner;
p. 88
1909 - Harpactocarcinus souverbiei A. Milne Edwards in Toniolo;
p. 293, PI. 26 (fig. 5)
1929 - Harpactocarcinus souverbiei A. Milne Edwards in Glaessner;
p. 206
1969 - Harpactoxanthopsis souverbiei (A. Milne Edwards) in Via
Boada; p. 300, 383
2000 - Harpactoxanthopsis souwerbiei (A. Milne Edwards) in Beschin,
De Angeli & Alberti; p. 1 5
2003 - Harpactoxanthopsis souwerbiei (A. Milne Edwards) in
Schweitzer; p. 1119
Holotype: MNHN R03823 (syntype).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Bittner (1875) reported this species
from Verona area (repository and catalogue number
unknown)
Note: this species is also reported from thè Eocene of
S France and Istria (Croazia).
Neozanthopsis Schweitzer, 2003
Type species: Harpactocarcinus americanus Rath-
bun, 1928.
Stratigraphic range: Eocene.
Neozanthopsis bruckmanni (v. Meyer, 1 862)
1859 - Xanthopsis hispidiformis in Reuss; p. 46, PI. 12 (fig. 3), PI. 13
(fig. 6), PI. 14 (figs. 1-4), PI. 23 (figs. 3-5)
1862 - Xanthopsis Bruckmanni v. Meyer; p. 152, PI. 16 (figs. 5-11),
PI. 17 (figs. 1.3)
1863 - Xanthopsis Bruckmanni v. Meyer in A. Milne Edwards; p. 322,
PI. 7 (figs. 3-4)
1863 - Xanthopsis Andreae Schafhàutl; p. 229, PI. 61 (figs. 2-3)
1863 - Xanthopsis Griintensis Schafhàutl; p. 228, PI. 62 (fig. 4)
1863 - Xanthopsis sonthofensis v. Meyer in Schafhàutl; p. 227, PI. 61
(figs. 5-6?)
1 877 - Xanthopsis sonthofensis v. Meyer in Mayer; p. 97
1925 -Xanthopsis Bruckmanni v. Meyer in Schlosser; p. 144
1929 - Xanthopsis Bruckmanni v. Meyer in Glaessner; p. 396
1930 - Xanthopsis Bruckmanni v. Meyer in Glaessner; p. 159
1959 - Xanthopsis Bruckmanni v. Meyer in Via Boada; p. 383
1969 - Xanthopsis Bruckmanni v. Meyer in Via Boada; p. 270, Text-
fig. 32, PI. 29 (figs. 1-3)
1973 -Xanthopsis bruckmanni v. Meyer in Via Boada; p. 62
1 989 - Zanthopsis bruckmanni (v. Meyer) in Solé & Via Boada; p. 32
2000 - Zanthopsis bruckmanni (v. Meyer) in Beschin, De Angeli &
Alberti; p. 14, PI. 1 (figs. 1-4)
2001 - Zanthopsis bruckmanni (v. Meyer) in De Angeli & Beschin;
p. 35, Text-fig. 30
2003 - Neozanthopsis bruchmanni (v. Meyer) in Schweitzer; p. 1119
2004 - Neozanthopsis bruckmanni (v. Meyer) in Beschin & De Angeli;
p. 20
Holotype: unknown.
Stratigraphic range: lower Eocene (Ypresian).
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
79
Occurrence: Veneto.
Material: Beschin et al. (2000) reported one speci¬
men (MCZ 1648) from “Rossi” quarry of Monte di Malo
(Vicenza).
Note: this species is also reported from thè Eocene of
Austria, Switzerland, Germany and Spain.
Superfamily Potamoidea Ortmann, 1 896
Family Potamidae Ortmann, 1896
Potamon Savigny, 1816
Type species: Potamon fluviatilis Savigny, 1816.
Stratigraphic range: upper Miocene - Recent.
1 Potamon castellinense (Szombathy, 1916)
187 4 - Pseudothelphusa speciosa Capellini; p. 39, PI. 7 (figs. 1-2)
1885 - Pseudothelphusa speciosa Capellini in Mercanti; p. 214, PI. 2
(fig. 15)
1904 - Pseudothelphusa speciosa Capellini in Capellini; p. 23 1
1916 - Pseudothelphusa speciosa Capellini in Rathbun; p. 415
1929 - Potamon ? castellinense (Szombathy) in Glaessner; p. 337
Stratigraphic range: Miocene.
Occurrence: Toscana.
Material: Capellini (1874) reported this species from
Valle del Marmolaio (Livorno) (repository and catalogue
number unknown).
Subsection Thoracotremata Guinot, 1977
Superfamily Pinnotheroidea De Haan, 1833
Family Pinnotheridae De Haan, 1833
Asthenognathus Stimpson, 1858
Type species: Asthenognatus inaequipes Stimpson,
1858, by originai designation.
Stratigraphic range: Oligocene - Recent.
Asthenognathus laverdensis De Angeli & Garassino,
2006
2006b - Asthenognathus laverdensis De Angeli & Garassino; p. 297,
Text-figs. 2-4
Holotype: MSNM Ì26744.
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli & Garassino (2006b) reported 15
specimens (MSNM from Ì26745 to Ì26752; MCZ from
2467 to 2472) from Laverda (Vicenza).
Superfamily Ocypodoidea Rafinesque-Schmaltz, 1815
Family Ocypodidae Rafinesque-Schmaltz, 1815
Archaeocypoda Secretan, 1975
Type species: Archaeocypoda veronensis Secretan,
1975.
Stratigraphic range: middle Eocene (Lutetian).
Archaeocypoda veronensis Secretan, 1975
1975 - Archaeocypoda veronensis Secretan; p. 363, Text-figs. 21-22,
Pls. 23 (2), 24, 25
Holotype: MSNM i45 (45bis).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Secretan (1975) reported three specimens
(MSNM i45, i46, 103) from Monte Bolca (Verona).
Family Palicidae Bouvier, 1898
Eopalicus Beschin, Busulini, De Angeli & Tessier, 1996
Type species: Eopalicus squamosus Beschin,
Busulini, De Angeli & Tessier, 1996.
Stratigraphic range: middle Eocene (Lutetian) -
lower Oligocene (Rupelian).
Eopalicus imbricatus De Angeli & Beschin, 2000
2000 - Eopalicus imbricatus De Angeli & Beschin; p. 9, Text-fig. 3
(2), PI. 1 (figs. 4-6)
2001 - Eopalicus imbricatus De Angeli & Beschin in De Angeli &
Beschin; p. 40, Text-fig. 34 (2)
2003 - Eopalicus imbricatus De Angeli & Beschin in Beschin &
De Angeli; p. 1 1
Holotype: MCZ 2034 (I.G. 286288).
Stratigraphic range: middle/upper Eocene (Barto-
nian - Priabonian).
Occurrence: Veneto.
Material: De Angeli & Beschin (2000) reported 24
specimens (MCZ from 2034 to 2057) from “Alonte”
quarry (Vicenza).
Eopalicus semicarinatus De Angeli & Beschin, 2000
2000 - Eopalicus semicarinatus De Angeli & Beschin; p. 10, Text-
fig. 3 (3), PI. 1 (fig. 3)
2001 - Eopalicus semicarinatus De Angeli & Beschin in De Angeli &
Beschin; p. 40, Text-fig. 34 (3)
2003 - Eopalicus semicarinatus De Angeli & Beschin in Beschin &
De Angeli; p. 1 1
Holotype: MCZ 2058 (I.G. 286312).
Stratigraphic range: lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: De Angeli & Beschin (2000) reported one
specimen (MCZ 2058) from Monte Lungo (Vicenza).
Eopalicus squamosus Beschin, Busulini, De Angeli &
Tessier, 1996
1996a - Eopalicus squamosus Beschin, Busulini, De Angeli &
Tessier; p. 77, Text-fig. 2, PI. 1 (figs. 1-3)
2000 - Eopalicus squamosus Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 8, Text-fig. 3 (1), PI. 1 (figs. 1-2)
200 1 - Eopalicus squamosus Beschin, Busulini, De Angeli & Tessier in
De Angeli & Beschin; p. 39, Text-fig. 34 (1)
2003 - Eopalicus squamosus Beschin, Busulini, De Angeli & Tessier in
Beschin & De Angeli; p. 1 1
80
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
2004 - Eopalicus squamosus Beschin, Busulini, De Angeli & Tessier in
Beschin, Busulini, De Angeli & Tessier; p. 115
Holotype: MCZ 1479 (I.G. 286354).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Beschin et al. (1996a) reported six speci-
mens (MCZ 1280, from 1479 to 1483) from “Main”
quarry of Arzignano (Vicenza) and “Albanello” quarry
of Nogarole Vicentino (Vicenza); De Angeli & Beschin
(2000) reported two specimens (MCZ 2032, 2033) from
“Main” quarry of Arzignano (Vicenza).
Spinipalicus Beschin & De Angeli, 2003
Type species: Spinipalicus italicus Beschin & De
Angeli, 2003.
Stratigraphic range: middle - upper Eocene (Lute¬
tian - Priabonian).
Spinipalicus italicus Beschin & De Angeli, 2003
2003 - Spinipalicus italicus Beschin & De Angeli; p. 9, Text-figs. 2-4
Holotype: MCZ 2390 (I.G. 296609).
Stratigraphic range: middle - upper Eocene (Lute¬
tian - Priabonian).
Occurrence: Veneto.
Material: Beschin & De Angeli (2003) reported
four specimens (MCZ 2390, 2391, 2392, 2393) from
“Lovara” quarry of Chiampo, Nanto, and Alonte quarry
(Vicenza).
Superfamily Grapsoidea MacLeay, 1838
Family Grapsidae MacLeay, 1838
Daragrapsus Miiller & Collins, 1991
Type species: Daragrapsus trispinosus Miiller & Col¬
lins, 1991.
Stratigraphic range: Eocene - Oligocene.
Daragrapsus trispinosus Miiller & Collins, 1 99 1
1991 - Daragrapsus trispinosus Miiller & Collins; p. 88, Text-fig. 5 h,
PI. 7(figs. 9-10, 12-14)
2001 - Daragrapsus trispinosus Miiller & Collins in Beschin,
De Angeli & Checchi; p. 28, Text-fig. 9, PI. 3 (figs. 5-6)
2001 - Daragrapsus trispinosus Miiller & Collins in De Angeli &
Beschin; p. 38
200 1 - Daragrapsus trispinosus Miiller & Collins in Karasawa & Kato;
p. 271,272
Holotype: MAFI EK-30.1 (M.9 1-209).
Stratigraphic range: upper Eocene (Priabonian) -
lower Oligocene (Rupelian).
Occurrence: Veneto.
Material: Beschin et al. (2001) reported eight speci¬
mens (MCZ 2066, 2067, 2071, 2072, 2073, 2074, 2117,
2124) from Castelgomberto (Vicenza).
Note: this species is also reported from thè upper
Eocene (Priabonian) of Hungary.
Daranyia Lòrenthey, 1 90 1
Type species: Daranyia granulata Lòrenthey, 1901.
Stratigraphic range: upper Eocene (Priabonian).
Daranyia fabianii Di Salvo, 1933
1933 - Daranyia Fabianii Di Salvo; p. 40, PI. 2 (figs. 5 a-d)
Holotype: MGUP 2620.
Stratigraphic range: upper Eocene (Priabonian).
Occurrence: Sicilia.
Material: Di Salvo (1933) reported one specimen
(MGUP 2620) from Bichinello (Palermo).
Grapsus Lamarck, 1801
Type species: Cancer grapsus Linnaeus, 1758.
Stratigraphic range: Miocene - Recent.
Grapsus sp.
I
1889 - Grapsus sp. in Ristori; p. 402, PI. 15 (figs. 2-3)
1974 - Grapsus sp. in Mastrorilli; p. 4
Stratigraphic range: Miocene.
Occurrence: Liguria.
Material: Ristori (1889) reported two specimens from
Sassello (Savona), housed in don Perrando’s collection,
today lost.
Pseudodaranyia Tessier, Beschin, Busulini & De Angeli,
1999
Type species: Pseudodaranyia carinata Tessier,
Beschin, Busulini & De Angeli, 1999.
Stratigraphic range: middle Eocene (Lutetian).
Pseudodaranyia carinata Tessier, Beschin, Busulini &
De Angeli, 1999
1999 - Pseudodaranyia carinata Tessier, Beschin, Busulini & De
Angeli; p. 99, Text-fig. 4, PI. 2 (figs. 4-5)
2001 - Pseudodaranyia carinata Tessier, Beschin, Busulini & De
Angeli in De Angeli & Beschin; p. 39
2004 - Pseudodaranyia carinata Tessier, Beschin, Busulini & De
Angeli in Beschin, Busulini, De Angeli & Tessier; p. 116
Holotype: MCZ 1599 (I.G. 284666).
Stratigraphic range: middle Eocene (Lutetian).
Occurrence: Veneto.
Material: Tessier et al. (1999) reported two speci¬
mens (MCZ 1599, 1600) from “Main” quarry of Arzi¬
gnano (Vicenza).
r
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
ACKNOWLEDGEMENTS
We wish to thank F. Barbagli, Museo Zoologico de
“La Specola” di Firenze, and F. Farsi, Accademia dei
Fisiocratici di Siena, for thè useful help to identify thè
originai specimens studied by Ristori, housed in thè
Accademia dei Fisiocratici di Siena; M. C. Bonci and
G. Vannucci, Dipartimento per lo Studio del Territorio e
delle sue Risorse (DIP.TE.RIS) di Genova, for thè useful
help to identify thè originai specimens of don Perran-
do’s collection, studied by Ristori; A. Ceregato, Palazzo
Poggi, Bologna, for permitting study of thè specimen of
Ranina aldrovandiv, D. Ormezzano, Museo Regionale di
Scienze Naturali di Torino, for thè useful help to identify
thè originai specimens studied by Crema; J.-M. Pacaud,
Muséum national d’Histoire naturalle, Paris, for thè
useful help to identify thè originai specimens studied by
A. Milne Èdwards; O. Schultz, Naturistorisches Museum,
Wien, for thè useful research of thè originai specimen
studied by Reuss; R. M. Feldmann and C. E. Schweitzer,
Geology Department, Kent State University (Ohio) for
thè useful help to identify thè types housed in Museum
fur Naturkunde der Humboldt-Universitàt, Berlin; P.
Artal, Seminario of Barcelona, for thè useful help to iden¬
tify some types of Via Boada’s collection. Moreover, we
wish to thank M. Fomasiero and L. Dal Favero, Museo
di Geologia e Paleontologia dell’Università di Padova,
B. Favaretto, Museo di Storia Naturale di Venezia, V.
Frisone, Museo Civico “G. Zannato” di Montecchio
Maggiore (Vicenza), and B. Pallozzi, Museo Civico
“D. Dal Lago” di Valdagno (Vicenza) to have given us
useful informations about thè palaeontological collec-
tions of their Museums. Finally, we wish to thank R. M.
Feldmann, Geology Department, Kent State University
(Ohio), for careful review and criticism.
82
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
BIBLIOGRAPHY
Airaghi C., 1 905 - Brachiuri nuovi o poco noti pel Terzia¬
rio Veneto. Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano,
Milano, 44: 202-209.
Allasinaz A., 1987 - Brachyura Decapoda oligocenici
(Rupeliano) del Bacino Ligure Piemontese. Bollettino
del Museo regionale di Scienze naturali di Torino,
Torino, 5 (2): 509-566.
Ancona L., 1966 - Esemplari di Ranina (Decapodi, Bra¬
chiuri) eccezionalmente ben conservati nell’Eocene
medio della Valle del Chiampo (Vicenza). Memo¬
rie del Museo Civico di Storia Naturale di Verona,
Verona, 14: 401-408.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1985 - Il genere Micromaia Bittner (Crustacea,
Decapoda) nel Terziario dell’area dei Berici e dei
Lessini, con descrizione di tre nuove specie. Lavori
- Società Veneziana di Scienze Naturali, Venezia, 10:
97-119.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1988 - Raninidae del Terziario berico-lessineo (Italia
settentrionale). Lavori - Società Veneziana di Scienze
Naturali, Venezia, 13: 155-215.
Beschin C., Busulini A., De Angeli A. & Tessier G., 1994
- 1 Crostacei eocenici della cava “Boschetto” di Noga-
role Vicentino (Vicenza - Italia settentrionale). Lavori
- Società Veneziana di Scienze Naturali, Venezia, 19:
159-215.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1 996a - Eopalicus nuovo genere di brachiuro (Deca¬
poda) del Terziario Veneto (Italia settentrionale).
Lavori - Società Veneziana di Scienze Naturali, Vene¬
zia, 21: 75-82.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1996b - Retroplumoidea (Crustacea, Brachyura) nel
Terziario del Vicentino (Italia settentrionale). Lavori
- Società Veneziana di Scienze Naturali, Venezia, 21:
83-102.
Beschin C., Busulini A., De Angeli A. & Tessier G., 2002
- Aggiornamento ai crostacei eocenici di cava “Main”
di Arzignano (Vicenza, Italia settentrionale) (Crusta¬
cea, Decapoda). Studi e Ricerche - Associazione Amici
Museo Zannato - Museo Civico “G. Zannato”, Mon-
tecchio Maggiore (Vicenza), 2002: 7-28.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
2004 - The Eocene decapod crustacean fauna of thè
“Main” quarry in Arzignano (Vicenza - NE Italy) with
thè description of a new species of Raninidae. Lavori
- Società Veneziana di Scienze Naturali, Venezia, 29:
109-117.
Beschin C., Busulini A., De Angeli A., Tessier G. &
Ungaro S., 1991 - Due nuovi generi di Raninidae del¬
l’Eocene del Veneto (Italia). Lavori - Società Vene¬
ziana di Scienze Naturali, Venezia, 16: 1 87-212.
Beschin C., Busulini A., De Angeli A., Tessier G. &
Ungaro S., 1998 - Crostacei eocenici di “Cava Rossi”
presso Monte di Malo (Vicenza - Italia settentrionale).
Studi Trentini di Scienze Naturali - Acta Geologica,
Trento, 73 (1996): 7-34.
Beschin C., Busulini A., De Angeli A., Tessier G. &
Ungaro S., 2000 - The fauna of thè Gecchelina quarry
at Monte di Malo (Vicenza - Northern Italy): a preli-
minary study. Extended abstracts - Studi e Ricerche
- Associazione Amici Museo Zannato - Museo Civico
“G. Zannato”, Montecchio Maggiore (Vicenza),
2000: 7-10.
Beschin C., Checchi A. & Ungaro S., 1996 - Crostacei
brachiuri dell’Oligocene di Castelgomberto (Lessini
orientali). Studi e Ricerche - Associazione Amici
Museo Zannato - Museo Civico ‘‘G. Zannato ”, Mon¬
tecchio Maggiore (Vicenza), 1996: 1 1-20.
Beschin C. & De Angeli A., 1984 - Nuove forme di
Anomura Hippidea: Albunea cuisiana sp. n. e Albunea
lutetiana sp. n. Lavori - Società Veneziana di Scienze
Naturali, Venezia, 9 (1): 93-105.
Beschin C. & De Angeli A., 2003 - Spinipalicus italicus,
nuovo genere e specie di Palicidae (Crustacea, Deca¬
poda) dell’Eocene del Vicentino (Italia settentrionale).
Studi e Ricerche - Associazione Amici Museo Zannato
- Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 2003: 7-12.
Beschin C. & De Angeli A., 2004 - Nuovi brachiuri eoce¬
nici dei Monti Lessini vicentini (Italia nordorientale).
Studi e Ricerche - Associazione Amici Museo Zannato
- Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 11: 13-22.
Beschin C., De Angeli A. & Alberti R., 2000 - Zan-
thopsis bruckmanni (Meyer) (Crustacea, Decapoda)
dell’Eocene del Vicentino (Italia settentrionale). Studi
e Ricerche - Associazione Amici Museo Zannato -
Museo Civico “G. Zannato”, Montecchio Maggiore
(Vicenza), 2000: 13-16.
Beschin C., De Angeli A. & Checchi A., 2001 - Crosta¬
cei decapodi associati a coralli della “Formazione di
Castelgomberto” (Oligocene) (Vicenza - Italia Set¬
tentrionale). Studi e Ricerche - Associazione Amici
Museo Zannato - Museo Civico “G. Zannato ”, Mon¬
tecchio Maggiore (Vicenza), 2001: 13-30.
Beschin C., De Angeli A., Checchi A. & Mietto P., 2006 -
Crostacei del Priaboniano di Priabona (Vicenza - Italia
settentrionale). Lavori - Società Veneziana di Scienze
Naturali, Venezia, 31: 95-1 12.
Beschin C., De Angeli A., Checchi A. & Zarantonello G., ,
2005 - Crostacei eocenici di Grola di Comedo Vicen¬
tino presso Spagnago (Vicenza, Italia Settentrionale).
Studi e Ricerche - Associazione Amici Museo Zannato
- Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 12: 5-35.
Beschin C., De Angeli A. & Garassino A., 2001 -Justitia
vicetina n. sp. (Crustacea, Decapoda) dell’Eocene di
Chiampo (Vicenza, Italia settentrionale). Studi Tren¬
tini di Scienze Naturali - Acta Geologica, Trento, 76
(1999): 89-97.
Beschin C. & Garassino A., 1999 - Penaeus vanzii n.
sp. e Penaeus sorbinii n. sp. (Crustacea, Decapoda)
dell’Oligocene della Valle del Ponte (Laverda) e Sal-
cedo (Vicenza, N. Italia). Atti della Società italiana
di Scienze naturali e del Museo civico di Storia natu¬
rale in Milano, Milano, 140 (2): 189-208.
Beschin C. & Santi L., 1997 - Cancer sismondai Meyer
(Crustacea, Decapoda) nelle argille plioceniche di
Vignola sul Panaro (Modena). Studi e Ricerche - Asso¬
ciazione Amici Museo Civico “G. Zannato”, Montec¬
chio Maggiore (Vicenza), 1997: 11-16.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
83
Bittner A., 1875 - Die Brachyuren des Vicentinischen
Tertiàrgebirges. Denkschriften Akademie Wissen¬
schaften , Wien, 34: 63-106.
Bittner A., 1883 - Neue Beitrage zur Kenntniss der
Brachyuren-Fauna des Altertiàrs von Vicenza und
Verona. Denkschriften Akademie Wissenschaften,
Wien, 46: 299-316.
Bittner A., 1884 - Beitrage zur Kenntniss tertiàrer Bra-
chyuren-Faunen. Denkschriften Akademie Wissen-
schaften, Wien, 48: 15-30.
Bittner A., 1886 - Neue Brachyuren des Eocaens von
Verona. Sitzungsberichte Akademie Wissenschaften ,
Wien, 94: 44-55.
Bittner A., 1895 - Uber zwei ungeniigend bekannte
brachyure Crustaceen des Vicentinischen Eocàns.
Sitzungsberichte Akademie Wissenschaften., Wien,
104: 247-253.
Boyko C. B., 2004 - A new genus of fossil sand crab
(Amonura: Albuneidae) from thè Oligocene of Italy.
Palaeontology, London, 47 (4): 933-936.
Bonfiglio L. & Donadeo G., 1982 - Cancer sismondai
Meyer nel Pliocene di Torre dell’Orso (Puglia). Atti
della Società italiana di Scienze naturali e del Museo
civico di Storia naturale in Milano, Milano, 123: 255-
296.
Bortoluzzi S., 2002 - Invertebrati del limite Plio-Plei-
stocene di Strongoli (KR): preparazione, studio e
valorizzazione. Università degli Studi di Padova
(unpublished degree).
Bravi S. & Garassino A., 1998a - New biostratigraphic
and palaeoecologic observations on thè “Plattenkalk”
of thè Lower Cretaceous (Albian) of Pietraroia (Bene-
vento, S Italy), and its decapod crustacean assem-
blage. Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale in Milano, Milano,
138: 119-171.
Bravi S. & Garassino A., 1998b - “Plattenkalk” of thè
Lower Cretaceous (Albian) of Petina, in thè Albumi
Mounts (Campania, S Italy) and its decapod crusta¬
cean assemblage. Atti della Società italiana di Scienze
naturali e del Museo civico di Storia naturale in
Milano, Milano, 138: 89-118.
Bravi S., Coppa M. G., Garassino A. & Patricelli R., 1999
- Palaeomon vesolensis n. sp. (Crustacea, Decapoda)
from thè Plattenkalk of Vesole Mount (Salerno, S
Italy). Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale in Milano, Milano,
140 (2): 141-169.
Busulini A., Tessier G., Beschin C. & De Angeli A., 2003
- Boschettia giampietroi, nuovo genere e specie di
Portunidae (Crustacea, Decapoda) dell’Eocene medio
della Valle del Chiampo (Vicenza, Italia settentrio¬
nale). Studi e Ricerche - Associazione Amici Museo
Zannato - Museo Civico “G. Zannato”, Montecchio
Maggiore (Vicenza), 2003: 13-18.
Busulini A., Tessier G. & Visentin M., 1982 - Brachyura
della cava Main (Arzignano) - Lessini orientali
(Vicenza) (Crustacea, Decapoda). Lavori - Società
Veneziana di Scienze Naturali, Venezia, 7: 75-84.
Busulini A., Tessier G. & Visentin M., 1984 - Titanocar-
cinus aculeatus nuova specie di brachiuro nell’Eocene
del Veneto (Crustacea, Decapoda). Lavori - Società
Veneziana di Scienze Naturali, Venezia, 9(1): 107-
117.
Busulini A., Tessier G., Visentin M., Beschin C., De
Angeli A. & Rossi A., 1983 - Nuovo contributo alla
conoscenza dei brachimi Eocenici di Cava Main (Arzi¬
gnano) - Lessini Orientali (Vicenza). Lavori - Società
Veneziana di Scienze Naturali, Venezia, 8: 55-73.
Capellini G., 1874 - La Formazione Gessosa di Castel¬
lina Marittima e i suoi fossili. Memorie dell’Accade¬
mia di Scienze Istituzionali di Bologna, Bologna, 3
(4): 1-60.
Catullo T. A., 1854 - Sui Crostacei fossili della calcaria
grossolana del veronese. «Lettera al Signor Professore
O. F. Naumann di Lipsia». Annuario Reale Istituto
Geologico Vienna.
Checchia-Rispoli G., 1903 - Sopra un crostaceo dei Tufi
calcarei Post-Pliocenici dei dintorni di Palermo. Bol¬
lettino della Società Geologica Italiana, Roma, 22:
488-492.
Checchia-Rispoli G., 1904 - L’ Harpactocarcinus pun-
ctulatus Desm. dell’Eocene di Peschici nel Monte
Gargano. Bollettino della Società Zoologica Italiana,
Roma 2 (5): 49-57.
Checchia-Rispoli G., 1905 - I crostacei dell’Eocene dei
dintorni di Monreale in provincia di Palermo. Gior¬
nale di Scienze Naturali ed Economiche di Palermo,
Palermo, 25: 309-325.
Checchia-Rispoli G., 1907 - Sopra un crostaceo dell’Eo¬
cene medio dei dintorni di Bagheria in Provincia di
Palermo. Bollettino della Società Geologica Italiana,
Roma, 26 (1): 25-28.
Checchia-Rispoli G., 1914 - “ Distefania ” nuovo genere
di Brachimi del Cenomaniano della Sicilia. Bollettino
della Società Zoologica Italiana, Roma, ser. 3, 3 (12):
173-184.
Collins J. S. H. & Dieni I., 1995 - New decapod crusta-
ceans from thè Cenomanian Rudist Limestones of NE
Italy. Bulletin of Mizunami Fossil Museum, Mizu-
nami, 22: 61-12.
Colosi G., 1921 - Un nuovo crostaceo fossile: “ Hete -
roglyphea paronaé”. Atti della Reale Accademia di
Scienze di Torino, Torino, 56: 79-82.
Comaschi Caria I., 1950 - Crostacei decapodi nel Mio¬
cene (Elveziano) di Bosa in Sardegna. Rendiconti del
Semimario della Facoltà di Scienze dell ’Univesità di
Cagliari, Cagliari, 20 (3-4): 324-327.
Comaschi Caria I., 1956 - I crostacei miocenici della
Sardegna. Bollettino del Servizio Geologico d’Italia,
Roma, 28 (1-2): 283-290.
Crema C., 1895 - Sopra alcuni decapodi terziarii del
Piemonte. Atti della Reale Accademia di Scienze di
Torino, Torino, 30: 664-681.
Dainelli G., 1915 - L’Eocene Friulano - Monografia geo¬
logica e paleontologica. Editrici le “ Memorie Geogra¬
fiche”, Firenze.
Dalla Vecchia F. M., 1991 - Note sulla stratigrafia,
sedimentologia e paleontologia della Dolomia di
Forni (Triassico superiore) nella Valle del Rio Seazza
(Preone, Friuli-Venezia Giulia). Gortania - Atti del
Museo Friulano di Storia Naturale, Udine, 12 (1990):
7-30.
Dalla Vecchia F. M., 1993 - Segnalazione di crosta¬
cei nell’Unità Fonte Santa (Triassico sup.) presso
Filettino (Lazio, Italia). Gortania - Atti del Museo
Friulano di Storia Naturale, Udine, 14 (1992):
59-69.
84
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
De Angeli A., 1995 - Crostacei dell’Eocene superiore
di “Fontanella” di Grancona (Vicenza - Italia set¬
tentrionale). Studi e Ricerche - Associazione Amici
Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 1995: 7-24.
De Angeli A., 1997 - Lysiosquilla messinae, nuova specie
di crostaceo stomatopode del Terziario di Vicenza
(Nord Italia). Studi e Ricerche - Associazione Amici
Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 1997: 23-26.
De Angeli A., 1998 - Gli Albuneidae (Crustacea, Hip-
poidea) del Terziario vicentino (Italia settentrionale).
Studi e Ricerche - Associazione Amici Museo Zannato
- Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 1998: 17-20.
De Angeli A. & Beschin C., 1998 - Ceronnectes, nuovo
genere di brachiuro (Crustacea, Decapoda) dell’Eo¬
cene di Ungheria e Italia. Lavori - Società Veneziana
di Scienze Naturali, Venezia, 23: 87-91.
De Angeli A. & Beschin C., 1999 - I crostacei Matuti-
nae (Brachyura, Calappidae) dell’Eocene del Veneto
(Italia settentrionale). Studi e Ricerche - Associazione
Amici Museo Zannato - Museo Civico “G. Zannato ”,
Montecchio Maggiore (Vicenza), 1999: 1 1-22.
De Angeli A. & Beschin C., 2000 - Due nuove specie
di Eopalicus (Decapoda, Palicidae) nel Terziario del
Veneto (Italia settentrionale). Studi e Ricerche - Asso¬
ciazione Amici Museo Zannato - Museo Civico “G.
Zannato”, Montecchio Maggiore (Vicenza), 2000:
7-12.
De Angeli A. & Beschin C., 2001 - I Crostacei fossili
del territorio Vicentino. Natura Vicentina, Vicenza,
5: 5-54.
De Angeli A. & Beschin C., 2002 - Branchi oplax albertii,
nuova specie di Goneplacidae (Crustacea, Decapoda)
dell’Eocene di cava “Main” di Arzignano (Vicenza
- Italia settentrionale). Lavori - Società Veneziana di
Scienze Naturali, Venezia, 27: 125-130.
De Angeli A. & Beschin C., 2004 - Nucilobus bericus
sp. nov., nuovo crostaceo Leucosiidae dell’Eocene
superiore del Veneto (Vicenza - Italia settentrionale).
Lavori - Società Veneziana di Scienze Naturali, Vene¬
zia, 29: 119-122.
De Angeli A., Beschin C. & Checchi A., 2005 - Una
nuova specie di Albuneidae Stimpson, 1858 dell’Eo¬
cene della Valle del Chiampo (Vicenza, NE Italia)
e considerazioni sulle altre forme note (Decapoda,
Anomura, Hippoidea). Lavori - Società Veneziana di
Scienze Naturali, Venezia, 30: 85-91.
De Angeli & Garassino A., 2002 - Galatheid, chirostylid
and porcellanid decapods (Crustacea, Decapoda, Ano-
mura) from thè Eocene and Oligocene of Vicenza (N
Italy). Memorie della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale di Milano,
Milano, 30 (3): 3-31.
De Angeli A. & Garassino A., 2003 - Bittnerilia, new
genus for Lambrus eocaenus Bittner, 1883 (Decapoda,
Brachyura, Calappidae) from thè middle Eocene of
Veneto (N Italy). Atti della Società italiana di Scienze
naturali e del Museo civico di Storia naturale in
Milano, Milano, 144(1): 13-22.
De Angeli A. & Garassino A., 2006a - New report of
decapod crustaceans from thè Mesozoic and Ceno-
zoic of Friuli-Venezia Giulia (NE Italy). Atti della
Società italiana di Scienze naturali e del Museo
civico di Storia naturale in Milano, Milano, 147 (2):
267-294.
De Angeli A. & Garassino A., 2006b - Asthenognathus
laverdensis n. sp. (Decapoda: Brachyura: Pinnotheri-
dae) from thè lower Oligocene of Laverda (Vicenza,
NE Italy). Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano ,
Milano, 147 (2): 295-304.
De Angeli A. & Marangon S., 1992 - Necronectes schaf-
feri Glaessner, nel Miocene della Sardegna (Italia).
Lavori - Società Veneziana di Scienze Naturali, Vene¬
zia, 17: 175-182.
De Angeli A. & Marangon S., 2001 - Paralbunea
galantensis, nuova specie di anomuro oligocenico
del Bacino Ligure-Piemontese (Italia settentrionale).
Studi Trentini Scienze Naturali - Acta Geologica,
Trento, 76(1999): 99-105.
De Angeli A. & Marangon S., 2003a - Decapod crusta¬
ceans from thè Oligocene of thè Ligure Piemontese
Basin, northem Italy. Contributions to Zoology, The
Hague, 72(2-3): 101-104.
De Angeli A. & Marangon S., 2003b - Contributo alla
conoscenza dei Decapodi oligocenici del Bacino
Ligure Piemontese (Italia settentrionale). Atti della
Società italiana di Scienze naturali e del Museo
civico di Storia naturale in Milano, Milano, 144 (2):
185-196.
De Angeli A. & Messina V., 1992 - Upogebia perarolen-
sis nuova specie di crostaceo del Terziario del Veneto.
Lavori - Società Veneziana di Scienze Naturali, Vene¬
zia, 17: 183-191.
De Angeli A. & Messina V., 1996 - Pseudosquilla
berica nuova specie di Stomatopoda del Terziario
Veneto (Italia Settentrionale). Studi e Ricerche
- Associazione Amici Museo Civico “G. Zannato ”,
Montecchio Maggiore (Vicenza), 1996: 5-10.
De Angeli A. & Messina V., 1997 - Galatea weinfurteri
Bachmayer, 1950 (Crustacea, Anomura) nell’Oli¬
gocene di Perarolo (Vicenza, Nord Italia). Studi e
Ricerche - Associazione Amici Museo Civico “G.
Zannato”, Montecchio Maggiore (Vicenza), 1997:
17-21.
De Angeli A. & Rossi A., 2006 - Crostacei oligocenici
di Perarolo (Vicenza - Italia settentrionale), con la
descrizione di una nuova specie di Mysida e di Iso-
poda. Lavori - Società Veneziana di Scienze Naturali,
Venezia, 31: 85-93.
De Gregorio A., 1884 - Nuovi Decapodi titonici. Natura¬
lista Siciliano, Palermo, 3: 134.
De Gregorio A., 1894 - Description des faunes tertiaires
de la Vénétie. Monographie des fossiles éocèniques
(Etage Parisien) de Mont Postale. Annales de Géolo-
gie et de Paleontologie, Palermo, 14: 1-55.
De Gregorio A., 1 895 - Note sur certains Crustacés (Bra-
chiures) éocéniques. (Avec un catalogue de tous les
Crustacés de la Vénetie cités par les Auteurs). Annales
de Géo/ogie et de Paléontologie, Palermo, 18: 1-22.
Delle Cave L., 1981 - Catalogue of type specimens in
thè invertebrate palaeontological collections of thè
Museum of Geology and Palaeontology of thè Uni¬
versity of Florence (Italy). Crustacea, Decapoda. Atti
della Società Toscana di Scienze Naturali, Memorie,
Pisa, serie A, 88: 43-50.
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM 1TALY
85
Delle Cave L., 1988a - Jaxea cf. nocturna (Crustacea,
Decapoda, Anomura) from thè Early Pliocene of
Toscana, Italy. Bollettino della Società Paleontolo¬
gica Italiana, Modena, 27 (1): 3-10.
Delle Cave L., 1988b - Monodaeus bortolottii, a new
species of Brachyura (Crustacea, Decapoda) from thè
Pliocene of Toscana (Italy). Bollettino della Società
Paleontologica Italiana, Modena, 27 (2): 123-127.
Desmarest A.-G., 1822 - Histoire Naturelle des Cru-
stacés fossiles. Les crustacés proprement dits. F.-G.
Levrault, Paris.
Di Salvo G., 1933 - I Crostacei del Terziario inferiore
della provincia di Palermo. Giornale Scienze Naturali
ed Economiche di Palermo, Palermo, 37: 1-44.
Fabiani R., 1908 - Paleontologia dei Colli Berici. Memo¬
rie di Matematica e di Fisica della Società Italiana
delle Scienze, Roma, ser. 3, 15: 45-248.
Fabiani R., 191 Oa — I crostacei terziarii del Vicentino.
Bollettino del Museo Civico di Vicenza, Vicenza, 1
(1): 1-40.
Fabiani R., 191 Ob — Sulle specie di Ranina finora note
ed in particolare sulla Ranina Aldrovandii. Atti della
Accademia Veneto Trentino Istriana, ser. 3, 3: 85-102.
Fabiani R., 1911 - Di una nuova specie di Phlyctenodes
( Phl . Dalpiazi ) dell’Oligocene dei Berici. Bollettino
del Museo Civico di Vicenza, Vicenza, 1: 41-45.
Fabiani R., 1915-11 Paleogene del Veneto. Memorie del¬
l’Istituto Geologico della Reale Università di Padova,
Padova, 3: 283-289.
Feldmann R. M., Bice K. L., Schweitzer Hopkins C.,
Salva E. W. & Pickford K., 1998 - Decapod crusta-
ceans from thè Eocene Castle Hayne Formation, North
Carolina: paleoceanographic implications. Journal of
Paleontology, Lawrence, Memoir48: 1-28.
Forster R., 1984 - Bàrenkrebse (Crustacea, Decapoda)
aus dem Cenoman des Libanon und dem Eozan Ita-
liens. Mitteilung Bayerische Staatlische Palàontologi-
sche historische Geologische, Miinchen, 24: 57-66.
Fucini A., 1910 - L ’Eriphia Cocchii Rist. Annali Univer¬
sità Toscana, Firenze, 30: 1-5.
Garassino A., 1996 - The family Erymidae Van Straelen,
1924 and thè superfamily Glypheoidea Zittel, 1885 in
thè Sinemurian of Osteno in Lombardia (Crustacea,
Decapoda). Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano,
Milano, 135: 333-373.
Garassino A., 1997 - The species Glyphea tonelloi n.
sp. (Crustacea, Decapoda) in thè Lower Cretaceous
(Upper Barremian-Aptian) of thè Valley of Comappo
River (Udine, NE Italy). Gortania - Atti del Museo
Friulano di Storia Naturale, Udine, 19: 85-93.
Garassino A., 1998 - New study on decapod crustaceans
of thè Lower Cretaceous (Barremian-Aptian) of thè
Valley of Comappo River (Udine, NE Italy). Gortania
- Atti del Museo Friulano di Storia Naturale, Udine,
20: 59-73.
Garassino A., 1999 - Nuova segnalazione di crostacei
decapodi nel Cretacico inferiore di Vemasso (Udine,
NE Italia). Gortania - Atti del Museo Friulano di
Storia Naturale, Udine, 21 : 61-64.
Garassino A. 2000a - Glyphea rigoi n. sp. (Crustacea,
Decapoda) of Dolomia di Forni (Norian, Upper Trias-
sic) of Gamia (Udine, NE Italy). Gortania - Atti del
Museo Friulano di Storia Naturale, Udine, 22: 59-64.
Garassino A., 2000b - Palinurus sp. (Crustacea, Deca¬
poda) del Cretacico inferiore (Barremiano-Aptiano)
della Valle del Torrente Comappo (Udine, NE Italia).
Gortania - Atti del Museo Friulano di Storia Naturale,
Udine, 22: 65-68.
Garassino A. & Bravi S., 2003 - Palaemon antonellae
n. sp. (Crustacea, Decapoda, Caridea) from thè Lower
Cretaceous “Platydolomite” of Profeti (Caserta, S
Italy). Journal of Paleontology, Lawrence, 77 (3):
589-592.
Garassino A. & De Angeli A., 2004a - Decapod crusta-
cean fauna from thè Pliocene and Pleistocene of Arda,
Stirane and Fiume Enzas (Piacenza, Parma and Reggio
Emilia Provinces, N. Italy). Atti della Società italiana
di Scienze naturali e del Museo civico di Storia natu¬
rale in Milano, Milano, 145 (1): 29-57.
Garassino A. & De Angeli A., 2004b - Parthenope angu-
lifrons Latreille, 1825, and Atelecyclus rotundatus
(Olivi, 1792) from thè Sicilian (upper Pleistocene) of
Favignana Island (Egadi Islands, Sicilia, S Italy). Atti
della Società italiana di Scienze naturali e del Museo
civico di Storia naturale in Milano, Milano, 145 (1):
19-28.
Garassino A., De Angeli A. & De Polli R., 2003 - Report
of Metanephrops Jenkins, 1972 (Crustacea, Deca¬
poda, Nephropidae) from thè upper Eocene of Gran-
cona (Vicenza, N Italy). Atti della Società italiana di
Scienze naturali e del Museo civico di Storia naturale
in Milano, Milano, 144 (2): 383-392.
Garassino A., De Angeli A., Gallo L. M. & Pasini G., 2004
- Brachyuran and anomuran fauna from thè Cenozoic
of Piemonte (NW Italy). Atti della Società italiana di
Scienze naturali e del Museo civico di Storia naturale
in Milano, Milano, 145 (2): 251-281.
Garassino A. & Fomaciari A., 2000 - Cancer sismondai
Meyer, 1843 (Crustacea, Decapoda) in thè Pleistocene
deposits of Fiume Enza (Parma, Italy). Extended
abstract - Studi e Ricerche - Associazione Amici
Museo - Museo Civico “G. Zannato”, Montecchio
Maggiore (Vicenza), 2000: 29-30.
Garassino A. & Gironi B., 2005 - Proeryon hartmanni (v.
Meyer, 1835) (Crustacea, Decapoda, Eryonoidea) and
Archaeopalinurus cfr. A. levis Pinna, 1974 (Crusta¬
cea, Decapoda, Palinuroidea) from thè Lower Jurassic
(Toarcian) of Cesana Brianza-Suello (Lecco, N Italy).
Atti della Società italiana di Scienze naturali e del
Museo civico di Storia naturale in Milano, Milano,
146(1): 53-68.
Garassino A. & Gironi B., 2006 - Coleia boboi n. sp.
(Crustacea, Decapoda, Coleiidae) from thè Upper
Triassic (Rhaetian) of Monte Verzegnis (Udine, NE
Italy). Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale in Milano, Milano,
147 (1): 93-102.
Garassino A. & Novati M., 2001 - Justitia desmaresti
(Massalongo, 1854) (Crustacea, Decapoda) from thè
Lutetian (Middle Eocene) of Monte Bolca (Verona, N
Italy). Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale in Milano, Milano,
141 (2): 251-268.
Garassino A. & Schweigert G., 2006 - The Upper Juras¬
sic Solnhofen decapod crustacean fauna: review of
thè types from old description. Part I. Infraorders
Astacidea, Thalassinidea and Palinura. Memorie
86
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
della Società italiana di Scienze Naturali e del Museo
civico di Storia Naturate di Milano , Milano, 34 (1):
1-64.
Garassino A. & Teruzzi G., 1 990 - The genus Aeger Mun-
ster, 1839 in thè Sinemurian of Osteno in Lombardia
(Crustacea, Decapoda). Atti della Società italiana di
Scienze naturali e del Museo civico di Storia naturale
in Milano, Milano, 131 (5): 105-136.
Garassino A. & Teruzzi G., 1993 - A new decapod cru-
stacean assemblage from thè Upper Triassic of Lom¬
bardia (N. Italy). Paleontologia Lombarda, Nuova
serie, Milano, 1: 3-27.
Garassino A. & Teruzzi G., 1995 - Decapod crustaceans
from thè Lower Cretaceous of Vemasso (Udine, NE
Italy). Gortania - Atti del Museo Friulano di Storia
Naturale, Udine, 16 (1994): 77-88.
Garassino A. & Teruzzi G., 2001 - I crostacei decapodi
del Toarciano (Giurassico inferiore) di Sogno (Ber¬
gamo, N. Italia). Atti della Società italiana di Scienze
naturali e del Museo civico di Storia naturale in
Milano, Milano, 141 (2): 187-197.
Garassino A., Teruzzi, G. & Dalla Vecchia F. M., 1996
- The macruran decapod crustaceans of thè Dolomia
di Forni (Norian, Upper Triassic) of Camia (Udine,
NE Italy). Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano,
Milano, 136: 15-60.
Gemmellaro G. G., 1869 - Studi paleontologici sulla
fauna del calcare a Terebratula janitor del Nord di
Sicilia. Palermo, Parte I. Stabilimento Tipografico
Lao.
Gemmellaro M., 1914 - Crostacei e pesci fossili del
“Piano Siciliano” dei dintorni di Palermo. Giornale di
Scienze Naturali ed Economiche di Palermo, Palermo,
30: 73-94.
Glaessner M. F., 1 929 - Fossilium catalogus. I: Ammalia.
Crustacea decapoda. J. F. Pompecky ed., Berlin.
Glaessner M. F., 1969 - Crustacea Decapoda. In: Treatise
on Invertebrate Paleontology. Arthropoda 4 (2), Moore
R. C., Geologica/ Society of America and University
of Kansas, Lawrence: R399-R533, R626-R628.
Karasawa H. & Kato H., 2001 - The systematic status of
thè genus Miosesarma Karasawa, 1989 with a phylo-
genetic analysis within thè family Grapsidae and a
review of fossil records (Crustacea: Decapoda: Bra-
chyura). Paleontologica l Research, 5 (4): 259-275.
Larghi C., 2002 - Mithracia oppionii sp. nov. (Crustacea,
Decapoda, Brachyura) from thè Eocene of Chiampo
(Vicenza, Italy). Bulletin of Mizunami Fossil Museum,
Mizunami, 29: 61-68.
Larghi C., 2003 - First record of Oligocene retroplumid
crab (Crustacea: Decapoda: Brachyura) from Italy.
Bulletin of Mizunami Fossil Museum, Mizunami, 30:
57-60.
Lòrenthey I., 1898 - Beitràge zur Decapodenfauna des
Ungarischen Tertiàrs. Termész Fiizetek, Budapest, 21:
1-133.
Lòrenthey 1., 1907 - Adatok Sardinia harmadidòszaki
ràk-faunàjàhoz. Mathematiche Es Természettudo-
manvi Kòzlemenyek, Budapest, 19 (2): 243-296.
Lòrenthey I., 1909 - Beitràge zur Tertiàren dekapoden-
fauna Sardiniens. Mathematischen und Naturwis-
senschaftlichen Berichte aus Ungarn, Budapest, 24:
202-257.
Lovisato D., 1894 - Avanzi di Squilla nel Miocene medio
di Sardegna. Rendiconti della Reale Accademia dei i
Lincei, Roma, 3 (4): 205-209.
Lovisato D., 1902 - Le specie fossili finora trovate nel
calcare compatto di Bonaria e di San Bartolomeo.
Tipo-Litografia Commerciale, Cagliari: 1-21.
Malaroda R. 1950 - Il Lattorfiano del Montecchio di
Custozza (Colli Berici). I. I macrofossili. Memorie del
Museo Civico di Storia Naturale di Verona, Verona,
2: 147-215.
Marangon S. & De Angeli A., 1997 - Cherpiocarcinus,
nuovo genere di brachiuro (Decapoda) dell’ Oligocene
del Bacino Ligure-Piemontese (Italia settentrionale).
Lavori - Società Veneziana di Scienze Naturali, Vene¬
zia, 22: 97-106.
Marras G. & Ventura G., 1991 - Crostacei decapodi
del miocene di Sassari (Sardegna nord-occidentale).
Bollettino della Società Sarda di Scienze Naturali,
Sassari, 28: 105-1 19.
Martin J. W. & Davis G. E., 2001 - An Updated Clas-
sification of thè Recent Crustacea. Naturai History
Museum of Los Angeles County, Sciences Series, Los
Angeles, 39: 1-123.
Maxia C., 1946 - Su alcuni crostacei dei dintorni di
Roma. Bollettino del Reale Ufficio Geologico d’Italia,
Roma, 69 (7): 129-150.
Meneghini G., 1857 - Paléontologie de File de Sardai-
gne. In: A Voyage en Sardaigne, Ed. La Marmora,
Torino.
r
Michelotti G., 1861 - Etude sur le miocène inferieur de
l’Italie septentrionale. Mémoire de la Société Hollan-
dais des Sciences, Paris.
Milne Edwards A., 1860 - Histoire des Crustacés
Podophthalmaires fossiles. Annales des Sciences
Naturelles (Zoologie et Botarne), 4ieme serie Zoolo¬
gie, Paris, 14(3): 129-293.
Milne Edwards A., 1861 - Remarques sur la faune carci-
nologique des Terrains quatemaires. L ’Institut, Paris,
29: 88.
Milne Edwards A., 1862-1865 - Monographies des cru¬
stacés de la famille cancériens. Annales des Sciences
Naturelles (Zoologie et Botarne), Paris, Serie 4,
Volume 18 (1862): 31-85; Volume 20 (1863): 273-
324; Serie 5, Volume 1 (1864): 31-88; Volume 31
(1865): 297-351.
Milne Edwards A., 1 872 - Note sur quelques Crustacés
fossiles appartenant aux genres Ranina et Galenop-
sis. Annales Sciences Gèologique , Paris, 3, art. 3:
2,8.
Milne Edwards H., 1850 - Formation nummulitique de
FEspagne. In: Histoire des progrés de la Géologie de
1834 a 1849. d’Archiac A. (ed.). Paris. 3: 1-304.
Miiller P., 1993 - Neogene decapod crustaceans from
Catalonia. Scripta Museum Seminario Barcinonensis,
Barcelona, 225: 1-39.
Miiller P. & Collins J. S. H., 1991 - Late Eocene coral-
associated decapods (Crustacea) from Hungary.
Contributions to Tertiary and Quaternary Geology,
Budapest, 28: 47-92.
Miinster G., von 1842 - Beschreibung drei neuer
Arten Crustaciten. Beitràge zur Petrefactenkunde, .
Bayreuth, 5.
Oppenheim P., 1901 - Die Priabonaschichten und ihre
Fauna. Paleontographica, Stuttgart, 47: 1-348.
I
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
87
Oppenheim P., 1903 - Ueber die Ueberkippung von
S. Orso, das Tertiàr des Tretto und Fauna wie
Stellung der Schioschichten. Zeitschrift Deut-
schen Geologischen Gesellschaft, Hannover, 55:
98-235.
Piccoli G. & Mocellin L. G., 1962 - Studi sulla macro¬
fauna priaboniana di Priabona. Memorie deli 'Istituto
Geologico e Mineralogico dell 'Università di Padova ,
Padova, 23: 3-120.
Pinna G., 1974 - I crostacei della fauna triassica di Cene
in Val Seriana (Bergamo). Memorie della Società ita¬
liana di Scienze naturali e del Museo civico di Storia
naturale di Milano, Milano, 21 (1): 5-34.
Pinna G., 1976 - 1 crostacei triassici dell’Alta Valvestino.
“Natura Bresciana’’ - Annali del Museo Civico di
Storia Naturale, Brescia, 13: 33-42.
Pinna G., 1968 - Gli Erionidei della nuova fauna sine-
muriana a crostacei decapodi di Osteno in Lombardia.
Atti della Società italiana di Scienze naturali e del
Museo civico di Storia naturale in Milano, Milano,
107: 93-134.
Pinna G., 1969 - Due nuovi esemplari di Coleia viallii
Pinna, del Sinemuriano inferiore di Osteno in Lombar¬
dia (Crustacea, Decapoda). Annali del Museo di Storia
naturale di Genova, Genova, 77: 626-632.
Reuss A., 1859 - Zur Kenntniss fossiler Krabben. Denk-
schriften der Akademie der Wissenschaften, Wien, 17:
1-90.
Ristori G., 1886 - I crostacei brachiuri e anomuri del
Pliocene italiano. Bollettino della Società Geologica
Italiana, Roma, 5: 93-129.
Ristori G., 1888 - Alcuni crostacei del Miocene medio
italiano. Atti della Società Toscana di Scienze Natu¬
rali, Pisa, 9: 212-219.
Ristori G., 1889 - Crostacei Piemontesi del Miocene
Inferiore. Bollettino della Società Geologica Italiana,
Roma, 7:397-413.
Ristori G., 189 la - Contributo alla fauna carcinologica
del Pliocene italiano. Atti della Società Toscana di
Scienze Naturali, Pisa, 1 1 : 3-18.
Ristori G., 1 891b — I crostacei fossili di Monte Mario.
Atti della Società Toscana di Scienze Naturali, Pisa,
11: 19-25.
Ristori G., 1892a - Note di Carcinologia pliocenica. Atti
della Società Toscana di Scienze Naturali, processi
verbali, Pisa, 8: 86-89.
Ristori G., 1892b - Resti di crostacei nel Pliocene
dell’Isola di Pianosa. Atti della Società Toscana di
Scienze Naturali, processi verbali, Pisa, 8: 90.
Ristori G., 1892c - I crostacei fossili di Chiavon. Atti
della Società Toscana di Scienze Naturali, processi
verbali, Pisa, 8: 160-163.
Ristori G., 1893 - Il Titanocarcinus raulinianus A. M.
Edw. negli strati nummulitici del Gargano. Atti della
Società Toscana di Scienze Naturali, Pisa, 7 (8): 212-
215.
Ristori G., 1896 - Crostacei neogenici di Sardegna e di
alcune altre località italiane. Bollettino della Società
Geolologica Italiana, Roma, 15 (4): 504-513.
Rizzotto D., 1998 - Nuovo esemplare di Cyrtorhina
globosa nell’Eocene medio della Valle del Chiampo
(Vicenza). Studi e Ricerche - Associazione Amici
Museo Zannata - Museo Civico “ G . Zannato ”, Mon-
tecchio Maggiore (Vicenza), 1998: 21-24.
Ronzani C., 1818 - Osservazione sopra un fossile del-
l’Aldrovandi chiamato Sepite. Memorie di Storia
Naturale, Bologna, 7: 76.
Schram F. R. & Mtìller H. G., 2004 - Catalog and biblio-
graphy of thè fossil and recent stomatopoda. Backhuys
publishers, Leiden.
Schweigert G., Garassino A., Hall R. L., Hauff R. B.
and Karasawa H., 2003 - The Lower Jurassic lobster
Uncina Quenstedt, 1851 (Crustacea: Decapoda: Asta-
cidea: Uncinidae). Stuttgarter Beitràge Naturkunde,
Stuttgart, Ser. B, 332: 1-20.
Schweitzer C. E., 2003 - Utility of proxy characters for
classifìcation of fossils: an example from thè fossil
Xanthoidea (Crustacea: Decapoda: Brachyura). Jour¬
nal ofPaleontology, Lawrence, 77 (6): 1 107-1128.
Schweitzer C. E., Feldmann R. M., Gonzalea-Barba G.
& Cosovic V., in press - New Decapoda (Anomura,
Brachyura) from thè Eocene Bateque and Tepetate
formations, Baja California Sur, Mexico. Bulletin of
Mizunami Fossil Museum, Mizunami.
Secretan S., 1975 - Les crustacés du Monte Bolca. Museo
civico Storia Naturale - Studi e Ricerche sui giaci¬
menti terziari di Bolca, Verona, 2: 315-346.
Sismonda A., 1839 -Notizie intorno a due fossili trovati nei
colli di S. Stefano Roero. Memorie della Reale Accade¬
mia di Scienze di Torino, Torino, ser. 2, 1: 90-95.
Sismonda E., 1846 - Descrizione dei pesci e dei crostacei
fossili nel Piemonte. Memorie della Reale Accademia
di Scienze di Torino, Torino, ser. 2, 10: 1-89.
Sismonda E., 1861 - Appendice alla descrizione dei pesci
e dei crostacei fossili. Memorie della Reale Accade¬
mia di Scienze di Torino, Torino, ser. 2, 19: 1-24.
Teruzzi G., 1990 - The genus Coleia Broderip, 1835
(Crustacea, Decapoda) in thè Sinemurian of Osteno in
Lombardia. Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano,
Milano, 131 (4): 85-104.
Tessier G., Beschin C., Busulini A. & De Angeli A.,
1999 - Nuovi brachiuri eocenici della cava “Main”
di Arzignano (Vicenza - Italia settentrionale). Lavori
- Società Veneziana di Scienze Naturali, Venezia, 24:
93-105.
Tessier G., Busulini A., Beschin C. & De Angeli A., 2004 -
Segnalazione di Cyrtorhina globosa Beschin, Busu¬
lini, De Angeli & Tessier, 1988 (Crustacea, Decapoda,
Brachyura) nell’Eocene di Zovo di Bolca (Verona,
Italia settentrionale). Studi e Ricerche - Associazione
Amici Museo Zannato - Museo Civico “G. Zannato’’,
Montecchio Maggiore (Vicenza), 1 1 (2004): 7-12.
Tettoni W., 1923 - Crostacei pliocenici dell’ Appennino
Modenese. Atti della Società Naturalisti e Matematici
di Modena, Modena, s. 6, 1 (53): 160-162.
Varola A., 1965 - Nota preliminare su di un giacimento
a Cancer sismondai Meyer nella Provincia di Lecce.
Atti della Società Toscana di Scienze Naturali, Pisa,
ser. A, 72 (1): 295-298.
Varola A., 1981 - Crostacei decapodi neogenici della
Provincia Salentina (Italia). Thalassia Salentina, 1 1 :
3-51.
Via Boada L., 1959 - Decàpodos fósiles del Eoceno
espanol. Boletin del Istituto Geològico y Minerò de
Espaha, Madrid, 70: 331-402.
Via Boada L., 1969 - Crustacéos Decàpodos del Eoceno
espanol. Pirineos, Barcelona, 91-94: 1-479.
88
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Vicariotto G. & Beschin C., 1994 - Galathea wein-
furteri Bachmayer nell’Oligocene dei Monti Berici
(Italia settentrionale) (Crustacea, Anomura). Studi
e Ricerche - Associazione Amici Museo Zannato -
Museo Civico “G. Zannato ”, Montecchio Maggiore
(Vicenza), 1994: 5-11.
Vicariotto M., 1997 - Nuovo contributo alla conoscenza
dei crostacei fossili della cava “Boschetto” di Noga-
role Vicentino (Vicenza, Nord Italia). Studi e Ricerche
- Associazione Amici Museo Zannato - Museo Civico
“G. Zannato”, Montecchio Maggiore (Vicenza),
1997:27-30.
Vinassa de Regny P. E., 1896 - Platycarcinus sismondai
del Museo Parmense e il Pa/aeocarpilius macrochei-
lus del Museo Pisano. Rivista Italiana di Paleontolo¬
gia , Roma, 2: 124-129.
Vinassa de Regny P. E., 1897 - Contribuzioni alla
conoscenza dei crostacei fossili italiani. Simonellia
quiricensis n. gen. n. sp. del Pliocene di S. Quirico
d’Orcia. Rivista Italiana di Paleontologia, Roma, 3
(5-6): 19-25.
Wright C. W. & Collins J. S. H., 1972 - British Creta-
ceou Crabs. Pa/aeonto/ogical Society Mononograph,
London: 1-114.
Antonio De Angeli - Associazione Amici del Museo Zannato, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: a.garassino@tin.it
Catalog and bibliography of thè fossil Stomatopoda and Decapoda from Italy
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo I
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
89
INDEX
Introduction . Pag. 3
Abbreviations for institutions holding speci-
mens . Pag. 3
Systematic catalog . Pag. 4
Stomatopoda Latreille, 1 8 1 7 . Pag. 4
Gonodactyloidea Giesbrecht, 1910 . Pag. 4
Pseudosquillidae Manning, 1997 . Pag. 4
Pseudosquilla Dana, 1852 . Pag. 4
P. berica De Angeli & Messina, 1996 . Pag. 4
Lysiosquilloidea Giesbrecht, 1910 . Pag. 4
Lysiosquillidae Giesbrecht, 1910 . Pag. 4
Lysiosquilla Dana, 1852 . Pag. 4
L. antiqua (Miinster, 1 842) . Pag. 4
L. messinai De Angeli, 1997 . Pag. 4
Squilloidea Latreille, 1802 . Pag. 4
Squillidae Latreille, 1802 . Pag. 4
Squilla Fabricius, 1787 . Pag. 4
S. miocenica Lovisato, 1894 . Pag. 5
Decapoda Latreille, 1 802 . Pag. 5
Dendrobranchiata Bate, 1888 . Pag. 5
Penaeoidea Rafinesque-Schmaltz, 1815 . Pag. 5
fAegeridae Burkenroad, 1963 . Pag. 5
Aeger Miinster, 1839 . Pag. 5
A. elongatus Garassino & Teruzzi, 1990 . Pag. 5
A.foersteri Garassino & Teruzzi, 1990 . Pag. 5
A. macropus Garassino & Teruzzi, 1990 . Pag. 5
A. muensteri Garassino & Teruzzi, 1990 . Pag. 5
A. robustus Garassino & Teruzzi, 1990 . Pag. 5
A. rostrospinatus Garassino & Teruzzi, 1990 .... Pag. 5
Penaeidae Rafinesque-Schmaltz, 1815 . Pag. 5
Antrimpos Miinster, 1839 . Pag. 5
A. noricus Pinna, 1974 . Pag. 5
Dusa Miinster, 1839 . Pag. 6
D. longipes (Pinna, 1974) . Pag. 6
Dusa cfr. D. longipes (Pinna, 1974) . Pag. 6
Longichela Garassino & Teruzzi, 1993 . Pag. 6
L. orobica Garassino & Teruzzi, 1 993 . Pag. 6
L. cfr. L. orobica Garassino & Teruzzi, 1993 .... Pag. 6
Micropenaeus Bravi & Garassino, 1998 . Pag. 6
M. tenuirostris Bravi & Garassino, 1998 . Pag. 6
Penaeus Fabricius, 1798 . Pag. 7
P. bolcensis Secretan, 1975 . Pag. 7
P. cornappensis Garassino, 1998 . Pag. 7
P obtusus Secretan, 1975 . Pag. 7
P. sorbinii Beschin & Garassino, 1999 . Pag. 7
P. vanzii Beschin & Garassino, 1 999 . Pag. 7
P. vernassensis Garassino, 1995 . Pag. 7
Pseudobombur Secretan, 1975 . Pag. 7
P. nummu/iticus Secretan, 1975 . Pag. 7
Satyrocaris Garassino & Teruzzi, 1994 . Pag. 7
S. cristatus (Garassino & Teruzzi, 1 993) . Pag. 7
Caridea Dana, 1852 . Pag. 8
Oplophoridae Dana, 1852 . Pag. 8
Tonellocaris Garassino, 1 998 . Pag. 8
T. brevirostrata Garassino, 1 998 . Pag. 8
Palaeomonidae Rafinesque-Schmaltz, 1815 .... Pag. 8
Alburnia Bravi & Garassino, 1998 . Pag. 8
A. petinensis Bravi & Garassino, 1998 . Pag. 8
Palaemon Weber, 1795 . Pag. 8
P. antonellae Garassino & Bravi, 2003 . Pag. 8
P. fabricii Michelotti, 1861 . Pag. 8
P. vesolensis Bravi, Coppa, Garassino & Patri-
celli, 1999 . Pag. 8
Indeterminate family . Pag. 9
Acanthinopus Pinna, 1974 . Pag. 9
A. gibbosus Pinna, 1974 . Pag. 9
Leiothorax Pinna, 1974 . Pag. 9
A. triasicus Pinna, 1974 . Pag. 9
Parvocaris Bravi & Garassino, 1998 . Pag. 9
A. samnitica Bravi & Garassino, 1998 . Pag. 9
Pinnacaris Garassino & Teruzzi, 1 993 . Pag. 9
P. dentata Garassino & Teruzzi, 1993 . Pag. 10
Astacidea Latreille, 1802 . Pag. 10
Glypheidae Zittel, 1885 . Pag. 10
Glyphea v. Meyer, 1835 . Pag. 10
G. rigoi Garassino, 2000 . Pag. 10
G. tonelloi Garassino, 1997 . Pag. 10
G. tricarinata Garassino, 1996 . Pag. 10
fMecochiridae Van Straelen, 1925 . Pag. 10
Mecochirus Germar, 1827 . Pag. 10
M. germari Garassino, 1996 . Pag. 10
Pseudoglyphea Oppel, 1861 . Pag. 10
P. ancylochelis (Woodward, 1863) . Pag. 11
P. gigantea Garassino & Teruzzi, 1993 . Pag. 1 1
P. paronae (Colosi, 1921) . Pag. 11
f Erymidae Van Straelen, 1 924 . Pag. 1 1
Eryma v. Meyer, 1 840 . Pag. 1 1
E. meyeri Garassino, 1996 . Pag. 11
1E. rinellincola De Gregorio, 1884 . Pag. 11
Pustulina Quenstedt, 1857 . Pag. 11
P sinemuriana (Garassino, 1996) . Pag. 11
Nephropidae Dana, 1852 . Pag. 11
Hoploparia McCoy, 1 849 . Pag. 1 1
Metanephrops Jenkins, 1972 . Pag. 12
Nephropsis Wood-Mason, 1873 . Pag. 12
IPlatychelidae Glaessner, 1969 . Pag. 12
Glaessnericaris Garassino & Teruzzi, 1993 . Pag. 12
G. dubia (Pinna, 1974) . Pag. 12
G. macrochela Garassino & Teruzzi, 1993 . Pag. 12
tUncinidae Beurlen, 1928 . Pag. 12
Uncina Quenstedt, 1851 . Pag. 12
U. alpina Schweigert, Garassino, Hall, Hauff &
Karasawa, 2003 . Pag. 12
Thalassinidea Latreille, 1831 . Pag. 13
Thalassinoidea Latreille, 1831 . Pag. 13
Thalassinidae Latreille, 1831 . Pag. 13
Thalassina Latreille, 1806 . Pag. 13
Callianassoidea Dana, 1852 . Pag. 13
90
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
Callianassidae Dana, 1852 . Pag. 13
Callianassa Leach, 1814 . Pag. 13
C. calaritana Ristori, 1896 . Pag. 13
C. canavarii Ristori, 1889 . Pag. 13
C. desmarestiana A. Milne Edwards, 1860 . Pag. 13
C. cfr. C. ferox Bittner, 1893 . Pag. 13
C. michelotti A. Milne Edwards, 1860 . Pag. 14
C. pedemontana Crema, 1 895 . Pag. 14
C. cfr. C. rakosiensis Lòrenthey, 1898 . Pag. 14
C. rovasendae Crema, 1895 . Pag. 14
C. sismondai A. Milne Edwards, 1860 . Pag. 14
C. subterranea (Montagli, 1 808) . Pag. 14
C. subterranea var. dentata Ristori, 1891 . Pag. 14
Corallianassa Manning, 1987 . Pag. 15
C. rigoi De Angeli & Garassino, 2006 . Pag. 15
Eoglypturus Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 15
E. grolensis Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 15
Eucalliax Manning &Felder, 1991 . Pag. 15
E. vicetina Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 15
Neocallichirus Sakai, 1988 . Pag. 15
N. allegranzii Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 15
N. borensis Beschin, De Angeli, Checchi &
Mietto, 2006 . Pag. 16
N. fonisi Beschin, Busulini, De Angeli & Tes¬
sier, 2002 . Pag. 16
Ctenochelidae Manning & Felder, 1991 . Pag. 16
Callianopsis de Saint Laurent, 1973 . Pag. 16
C. microspineus Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 16
Ctenocheles Kishinouye, 1926 . Pag. 16
C. ornatus Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 16
C. valdellae (Fabiani, 1908) . Pag. 16
Laomediidae Borradaile, 1903 . Pag. 17
Jaxea Nardo, 1 847 . Pag. 17
J. cfr. J. nocturna Nardo, 1847 . Pag. 17
Upogebiidae Borradaile, 1903 . Pag. 17
Upogebia Leach, 1814 . Pag. 17
U. perarolensis De Angeli & Messina, 1992 .... Pag. 17
U. cfr. U. stellata (Montagu, 1808) . Pag. 17
Axiidae Huxley, 1879 . Pag. 17
Etallonia Oppel, 1861 . Pag. 17
Huxleycaris Bravi & Garassino, 1 998 . Pag. 1 7
H. beneventana Bravi & Garassino, 1998 . Pag. 17
Protaxius Beurlen, 1930 . Pag. 17
P. eocenicus Secretan, 1975 . Pag. 17
Indeterminate family . Pag. 18
Orhomalus Etallon, 1861 . Pag. 18
O. rotulensis De Gregorio, 1884 . Pag. 18
Palinura Latreille, 1802 . Pag. 18
fColeiidae Van Straelen, 1924 . Pag. 18
Coleia Broderip, 1835 . Pag. 18
C. boboi Garassino & Gironi, 2006 .
C. mediterranea Pinna, 1968 .
C. pianai Teruzzi, 1990 .
C. popeyei Teruzzi, 1 990 .
C. viallii Pinna, 1968 .
Proeryon Beurlen, 1928 .
P. hartmanni (v. Meyer, 1835) .
Pseudoco/eia Garassino & Teruzzi, 1993 .
P. mazzolenii Garassino & Teruzzi, 1 993 .
tEryonidae De Haan, 1841 .
Rosenfeldia Garassino, Teruzzi & Dalla Vec¬
chia, 1996 .
R. triasica Garassino, Teruzzi & Dalla Vecchia,
1996 .
Palinuridae Latreille, 1802 .
Archaeopalinurus Pinna, 1974 .
A. levis Pinna, 1974 .
Justitia Holthuis, 1946 .
J. desmaresti (Massalongo, 1854) .
J. vicetina Beschin, De Angeli & Garassino,
2001 .
Palinurus Weber, 1795 .
Scyllaridae Latreille, 1825 .
Parribacus Dana, 1 852 .
P cristatus Forster, 1984 .
Anomura MacLeay, 1838 .
Chirostylidae Ortmann, 1892 .
Eumunida Smith, 1883 .
E. pentacantha (Miiller & Collins, 1991) .
Galatheidae Samouelle, 1819 .
Calteagalathea De Angeli & Garassino, 2006 ..
C. friulana De Angeli & Garassino, 2006 .
Galathea Fabricius, 1793 .
G. affnis Ristori, 1886 .
G. berica De Angeli & Garassino, 2002 .
G. valmaranensis De Angeli & Garassino,
2002 .
G. cfr. G. weinfurteri Bachmayer, 1950 .
Acanthogalathea Miiller & Collins, 1991 .
A. feldmanni De Angeli & Garassino, 2002 .
A. parva (Miiller & Collins, 1991) .
Lessiniga/athea De Angeli & Garassino, 2002 .
L. regale De Angeli & Garassino, 2002 .
Palaeomunida Lòrenthey, 1901 .
P. defecta Lòrenthey, 1901 .
P. multicristata De Angeli & Garassino, 2002 ..
Spathagalathea De Angeli & Garassino, 2002 .
S. minuta De Angeli & Garassino, 2002 .
Porcellanidae Haworth, 1825 .
Beripetrolisthes De Angeli & Garassino, 2002 .
B. mulleri De Angeli & Garassino, 2002 .
Eopetrolisthes De Angeli & Garassino, 2002 ....
E. striatissimus (Miiller & Collins, 1991) .
Lobipetrolisthes De Angeli & Garassino, 2002 .
L. blowi De Angeli & Garassino, 2002 .
Longoporcellana Miiller & Collins, 1991 .
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
91
L. lobata De Angeli & Garassino, 2002 . Pag. 23
Petrolisthes Stimpson, 1858 . Pag. 23
P. bittneri De Angeli & Garassino, 2002 . Pag. 23
P vicetinus Beschin, De Angeli & Checchi, 200 1 Pag. 23
Pisidia Leach, 1 820 . Pag. 24
P. dorsosinuosa De Angeli & Garassino, 2002 . Pag. 24
Hippoidea Latreille, 1825 . Pag. 24
Albuneidae Stimpson, 1858 . Pag. 24
Albunea Weber, 1795 . Pag. 24
A. cuisiana Beschin & De Angeli, 1984 . Pag. 24
ltalialbunea Boyko, 2002 . Pag. 24
I. lutetiana (Beschin & De Angeli, 1984) . Pag. 24
Stemonopa Efford & Haig, 1968 . Pag. 24
S. prisca De Angeli, Beschin & Checchi, 2005 . Pag. 24
Zygopa Holthuis, 1961 . Pag. 24
Z. galantensis (De Angeli & Marangon, 2001) . Pag. 24
Paguroidea Latreille, 1802 . . Pag. 25
Diogenidae Ortmann, 1 892 . Pag. 25
Calcinus Dana, 1851 . Pag. 25
C. agnoensis Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 25
Dardanus Paulson, 1875 . Pag. 25
D. mediterraneus (Lòrenthey, 1909) . Pag. 25
Diogenes Dana, 1851 . Pag. 25
Eocalcinus Via Boada, 1959 . Pag. 25
E. cavus Beschin, Busulini, De Angeli & Tes-
sier, 2002 . Pag. 25
E. eocenicus Via Boada, 1959 . Pag. 26
Paguristes Dana, 1851 . Pag. 26
P. lineatuberculatus Beschin, De Angeli, Chec¬
chi & Mietto, 2006 . Pag. 26
P. prealpinus Beschin, De Angeli, Checchi & Pag. 26
Zarantonello, 2005 .
Petrochirus Stimpson, 1859 . Pag. 26
P. mezi (Lòrenthey, 1909) . Pag. 26
P. poscolensis Beschin, De Angeli, Checchi &
Mietto, 2006 . Pag. 26
Paguridae Latreille, 1802 . Pag. 26
Anapagurus Henderson, 1888 . Pag. 26
Pagurus Fabricius, 1775 . Pag. 26
P. arrosor (Herbst, 1794) . Pag. 27
P. manzonii (Ristori, 1888) . Pag. 27
P. mediterraneus Lòrenthey, 1909 . Pag. 27
P. squamosus Ristori, 1886 . Pag. 27
P. substriatus A. Milne Edwards, 1861 . Pag. 27
P. cfr. P. substriatus A. Milne Edwards, 1861 ... Pag. 27
Brachyura Latreille, 1 802 . Pag. 28
Podotremata Guinot, 1977 . Pag. 28
Dromiacea De Haan, 1833 . Pag. 28
Homolodromioidea Alcock, 1899 . Pag. 28
fProsopidae v. Meyer, 1860 . Pag. 28
Pithonoton v. Meyer, 1 842 . Pag. 28
P bidentatum (Reuss, 1858) . Pag. 28
P. grande (v. Meyer, 1860) . Pag. 28
P. marginatum (v. Meyer, 1 842) . Pag. 28
Indeterminate family . Pag. 28
Oxythyreus Reuss, 1858 . Pag. 28
Oxythyreus gibbosus Reuss, 1858 . Pag. 28
Dromioidea De Haan, 1833 . Pag. 29
Dromiidae De Haan, 1833 . Pag. 29
Dromia Weber, 1795 . Pag. 29
D. vulgaris H. Milne Edwards, 1837 . Pag. 29
Noetlingia Beurlen, 1928 . Pag. 29
N. claudiopolitana (Bittner, 1893) . Pag. 29
N. veronensis (Bittner, 1886) . Pag. 29
Dynomenidae Ortmann, 1892 . Pag. 29
Basinotopus McCoy, 1849 . Pag. 29
B. lamarcki (Desmarest, 1822) . Pag. 29
Cyamocarcinus Bittner, 1883 . Pag. 30
C. angustifrons Bittner, 1883 . Pag. 30
Cyclothyreus Remes, 1895 . Pag. 30
C. oxythyreiforme (G. G. Gemmellaro, 1869) ... Pag. 30
C. reussi (G. G. Gemmellaro, 1869) . Pag. 30
C. tithonium (G. G. Gemmellaro, 1869) . Pag. 30
Dromilites H. Milne Edwards, 1837 . Pag. 31
D. corvini (Bittner, 1893) . Pag. 31
D. hilarionis (Bittner, 1883) . Pag. 31
D. pastoris Via Boada, 1959 . Pag. 31
Dynomene Desmarest, 1 823 . Pag. 3 1
D. lessinea Beschin, De Angeli & Checchi,
2001 . Pag. 31
Gemmellarocarcinus Checchia-Rispoli, 1905 ... Pag. 31
G. lòrentheyi Checchia-Rispoli, 1905 . Pag. 31
Graptocarcinus Roemer, 1887 . Pag. 32
G. bellonii Collins & Dieni, 1995 . Pag. 32
Kromtitis Muller, 1984 . Pag. 32
K. tetratuberculatus Beschin, Busulini, De
Angeli & Tessier, 2002 . Pag. 32
Palaeodromites A. Milne Edwards, 1865 . Pag. 32
P. himeraensis (Checchia-Rispoli, 1914) . Pag. 32
Archaeobrachyura Guinot, 1977 . . Pag. 32
Raninoidea De Haan, 1839 . Pag. 32
Raninidae De Haan, 1839 . Pag. 32
Cosmonotus Adams & Withe, 1848 . Pag. 32
C. eocaenicus Beschin, Busulini, De Angeli &
Tessier, 1988 . Pag. 32
Cyrtorhina Monod, 1956 . Pag. 32
C. globosa Beschin, Busulini, De Angeli &
Tessier, 1988 . Pag. 32
C. oblonga Beschin, Busulini, De Angeli &
Tessier, 1988 . Pag. 33
Lianira Beschin, Busulini, De Angeli, Tessier &
Ungaro, 1991 . Pag. 33
L. beschini Beschin, Busulini, De Angeli, Tes¬
sier & Ungaro, 1991 . Pag. 33
L. convexa Beschin, Busulini, De Angeli, Tes¬
sier & Ungaro, 1991 . Pag. 33
Lophoranina Fabiani, 1910 . Pag. 33
L. aldrovandii Ronzani, 1918 . Pag. 33
L. bittneri (Lòrenthey, 1902) . Pag. 34
L. laevifrons (Bittner, 1875) . Pag. 34
L. marestiana (Kònig, 1825) . Pag. 34
92
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
L. marestiana var. avesana (Bittner, 1883) . Pag. 35
L. maxima Beschin, Busulini, De Angeli & Pag. 35
Tessier, 2004 .
L. reussi (Woodward, 1866) . Pag. 35
L. straeleni Via Boada, 1959 . Pag. 36
Lovarina Beschin, Busulini, De Angeli, Tessier Pag. 36
& Ungaro, 1991 .
L. cristata Beschin, Busulini, De Angeli, Tessier Pag. 36
& Ungaro, 1991 .
Lysirude Goeke, 1985 . Pag. 36
L. paronae (Crema, 1895) . Pag. 36
Notopoides Henderson, 1888 . Pag. 36
N. exiguus Beschin, Busulini, De Angeli &
Tessier, 1988 . Pag. 36
Notopus De Haan, 1841 . Pag. 36
N. beyrichi Bittner, 1875 . Pag. 36
Quasilaeviranina Tucker, 1998 . Pag. 37
Q. arzignanensis (Beschin, Busulini, De Angeli
& Tessier, 1988) . Pag. 37
Q. ombonii (Fabiani, 1910) . Pag. 37
Q. simplicìssima (Bittner, 1883) . Pag. 37
Ranilia H. Milne Edwards, 1837 . Pag. 37
R. punctulata Beschin, Busulini, De Angeli &
Tessier, 1988 . Pag. 37
Ranina Lamarck, 1801 . Pag. 38
R. bouilleana A. Milne Edwards, 1872 . Pag. 38
R. palmea E. Sismonda, 1846 . Pag. 38
R. propìnqua Ristori, 1891 . Pag. 38
R. speciosa (Miinster, 1840) . Pag. 38
Raninoides H. Milne Edwards, 1837 . Pag. 38
R. budapestiniensis (Lòrenthey, 1897) . Pag. 38
R. fabianii (Lòrenthey & Beurlen, 1929) . Pag. 39
R. cfr. R. fabianii (Lòrenthey & Beurlen, 1929) Pag. 39
R. notopoides (Bittner, 1883) . Pag. 39
R. pulchra (Beschin, Busulini, De Angeli &
Tessier, 1988) . Pag. 39
Tribolocephalus Ristori, 1886 . Pag. 39
T. laevis Ristori, 1886 . Pag. 39
Eubrachyura de Saint Laurent, 1980 . Pag. 39
Heterotremata Guinot, 1977 . Pag. 39
Dorippoidea MacLeay, 1838 . Pag. 39
Dorippidae MacLeay, 1838 . Pag. 39
Medorippe Manning & Holthuis, 1981 . Pag. 39
M. ampia Garassino, De Angeli, Gallo & Pasini,
2004 . Pag. 40
M. lanata (Linnaeus, 1767) . Pag. 40
fOrithopsidae Schweitzer, Feldmann, Fam,
Hessin, Hetrick, Nyborg & Ross, 2003 . Pag. 40
Cherpiocarcinus Marangon & De Angeli, 1997 Pag. 40
C. rostratus Marangon & De Angeli, 1997 . Pag. 40
Calappoidea H. Milne Edwards, 1837 . Pag. 40
Calappidae H. Milne Edwards, 1837 . Pag. 40
Bittnerilia De Angeli & Garassino, 2003 . Pag. 40
B. dentata Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 40
B. eocaena (Bittner, 1883) . Pag. 40
Calappa Weber, 1795 . Pag. 40
C. granulata (Linnaeus, 1758) . Pag. 40
Calappilia A. Milne Edwards, 1873 . Pag. 41
C. dacica Bittner, 1893 . Pag. 41
C. gemmata Beschin, Busulini, De Angeli &
Tessier, 1994 . Pag. 42
C. incisa Bittner, 1886 . Pag. 42
C. mainii Allasinaz, 1987 . Pag. 42
C. subovata Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 42
C. verrucosa A.. Milne Edwards, 1873 . Pag. 42
C. vicetina Fabiani, 1910 . Pag. 42
Mursiopsis Ristori, 1889 . Pag. 43
M. pustulosus Ristori, 1889 . Pag. 43
Hepatidae Stimpson, 1871 . Pag. 43
Hepatiscus Bittner, 1875 . Pag. 43
IH. distefanoi Checchia-Rispoli, 1905 . Pag. 43
H. minimus Beschin, Busulini, De Angeli &
Tessier, 1994 . Pag. 43
H. neumayri Bittner, 1875 . Pag. 43
H. pulchellus Bittner, 1875 . Pag. 43
Mainhepatiscus De Angeli & Beschin, 1999 .... Pag. 44
M. zannatoi De Angeli & Beschin, 1999 . Pag. 44
Osachila Stimpson, 1871 . Pag. 44
O. berica De Angeli & Beschin, 1999 . Pag. 44
Priabonella Beschin, De Angeli, Checchi &
Mietto, 2006 . Pag. 44
P. violatii Beschin, De Angeli, Checchi &
Mietto, 2006 . Pag. 44
Pseudohepatiscus Blow & Manning, 1996 . Pag. 44
P. silvanoi De Angeli & Beschin, 1999 . Pag. 44
Leucosioidea Samouelle, 1819 . Pag. 45
Leucosiidae Samouelle, 1819 . Pag. 45
Ebalia Leach, 1817 . Pag. 45
E. cranchii Leach, 1817 . Pag. 45
E. fucinii Ristori, 1 892 . Pag. 45
E. lamarmorai Lòrenthey, 1909 . Pag. 45
E. tuberosa Pennant, 1777 . Pag. 45
Hepatinulus Ristori, 1886 . Pag. 45
H. lovisatoi Lòrenthey, 1909 . Pag. 45
H. seguentiae Ristori, 1886 . Pag. 46
Ilia Leach, 1817 . Pag. 46
I. cfr. I. nucleus (Linnaeus, 1758) . Pag. 46
I. pliocaenica Ristori, 1891 . Pag. 46
Nucilobus Morris & Collins, 1991 . Pag. 46
N. bericus De Angeli & Beschin, 2004 . Pag. 46
Palaeomyra A. Milne Edwards in E. Sismonda,
1861 . Pag. 46
P. bispinosa A. Milne Edwards in E. Sismonda,
1861 . Pag. 46
Typilobus Stoliczka, 1871 . Pag. 46
T. semseyanus Lòrenthey, 1 897 . Pag. 46
Majoidea Samouelle, 1819 . Pag. 47
Majidae Samouelle, 1819 . Pag. 47
Maja Lamarck, 1801 . Pag. 47
M. miocaenica Lòrenthey, 1907 . Pag. 47
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
93
M. squinado (Herbst, 1788) . Pag. 47
Micromaia Bittner, 1875 . Pag. 47
M. elegans Beschin, Busulini, De Angeli &
Tessier, 1985 . Pag. 47
M. mainensis Beschin, Busulini, De Angeli &
Tessier, 1985 . Pag. 47
M. margaritata Fabiani, 1910 . Pag. 48
M. meneguzzoi Beschin, Busulini, De Angeli &
Tessier, 1985 . Pag. 48
M. priabonensis Oppenheim, 1901 . Pag. 48
M. tuberculata Bittner, 1875 . Pag. 48
Periacanthus Bittner, 1875 . Pag. 49
P. dallagoi Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 49
P. horridus Bittner, 1875 . Pag. 49
Mithracidae Balss, 1929 . Pag. 49
Micippa Leach, 1817 . Pag. 49
M. antiqua Beschin, De Angeli & Checchi,
2001 . Pag. 49
Mithracia Bell, 1858 . Pag. 49
M. margaritifera Beschin, Busulini, De Angeli
& Tessier, 1994 . Pag. 50
M. oppionii Larghi, 2002 . Pag. 50
Pisidae Dana, 1851 . Pag. 50
Pisa Leach, 1814 . Pag. 50
P. armata (Latreille, 1803) . Pag. 50
Parthenopoidea MacLeay, 1838 . Pag. 50
Dairidae Ng & Rodriguez, 1986 . Pag. 50
Daira De Haan, 1833 . Pag. 50
D. coronata Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag. 50
D. depressa (A. Milne Edwards, 1865) . Pag. 50
D. eocenica var. sicula (Di Salvo, 1933) . Pag. 50
D. salebrosa Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 51
Daldorfìidae Ng & Rodriguez, 1986 . Pag. 51
Daldorfìa Rathbun, 1 904 . Pag. 5 1
D. fabianii Beschin, De Angeli & Checchi,
2001 . Pag. 51
Parthenopidae MacLeay, 1838 . Pag. 51
Parthenope Weber, 1795 . Pag. 51
P. angulifrons Latreille, 1 825 . Pag. 5 1
P. nummuìitica (Bittner, 1875) . Pag. 51
Retroplumoidea Gill, 1894 . Pag. 52
Retroplumidae Gill, 1894 . Pag. 52
Loerenthopluma Beschin, Busulini, De Angeli
& Tessier, 1996 . Pag. 52
L. lata Beschin, Busulini, De Angeli & Tessier,
1996 . Pag. 52
Retrocypoda Via Boada, 1959 . Pag. 52
R. almelai Via Boada, 1959 . Pag. 52
Retropluma Gill, 1 894 . Pag. 52
R. craverii (Crema, 1895) . Pag. 52
R. eocenica Via Boada, 1959 . Pag. 52
R. cfr. R. eocenica Via Boada, 1959 . Pag. 53
Cancroidea Latreille, 1802 . Pag. 53
Atelecyclidae Ortmann, 1 893 . Pag. 53
Atelecyclus Leach, 1814 . Pag. 53
A. elegans Ristori, 1896 . Pag. 53
A. rotundatus (Olivi, 1792) . Pag. 53
Cancridae Latreille, 1802 . Pag. 53
Cancer Linnaeus, 1758 . Pag. 53
C. spinosus (Ristori, 1886) . Pag. 53
Ceronnectes De Angeli & Beschin, 1998 . Pag. 53
C. boeckhi (Lòrenthey, 1897) . Pag. 53
Lobocarcinus Reuss, 1857 . Pag. 54
L. sismondai (v. Meyer, 1843) . Pag. 54
Cheiragonidae Ortmann, 1893 . Pag. 55
Montezumella Rathbun, 1930 . Pag. 55
M. elegans (Lòrenthey & Beurlen, 1929) . Pag. 55
M. pumicosa Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 55
M. scabra Quayle & Collins, 1981 . Pag. 55
Portunoidea Rafinesque-Schmaltz, 1 8 1 5 . Pag. 55
Geryonidae Colosi, 1924 . Pag. 55
Coeloma A. Milne Edwards, 1865 . Pag. 55
C. isseli Ristori, 1886 . Pag. 56
C. sabatium Ristori, 1886 . Pag. 56
C. vigil A. Milne Edwards, 1865 . Pag. 56
Portunidae Rafinesque-Schmaltz, 1815 . Pag. 56
Boschettia Busulini, Tessier, Beschin & De
Angeli, 2003 . Pag. 56
B. giampietroi Busulini, Tessier, Beschin & De
Angeli, 2003 . Pag. 56
Carcinus Leach, 1814 . Pag. 56
Charybdis De Haan, 1833 . Pag. 57
C. antiqua (A. Milne Edwards, 1860) . Pag. 57
Enoplonotus A. Milne Edwards, 1860 . Pag. 57
E. armatus A. Milne Edwards, 1860 . Pag. 57
Eocharybdis Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 57
E. cristata Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 57
Liocarcinus Stimpson, 1871 . Pag. 57
L. cfr. L. rakosensis (Lòrenthey & Beurlen,
1929) . Pag. 57
Macropipus Prestandrea, 1833 . Pag. 57
M. ovalipes Secretan, 1975 . Pag. 57
Necronectes A. Milne Edwards, 1881 . Pag. 57
N. schafferi Glaessner, 1928 . Pag. 57
Neptocarcinus Lòrenthey, 1 897 . Pag. 58
N. millenaris Lòrenthey, 1897 . Pag. 58
Nogarolia Beschin, Busulini, De Angeli & Tes¬
sier, 1994 . Pag. 58
N. mirabilis Beschin, Busulini, De Angeli &
Tessier, 1994 . Pag. 58
Portunites Bell, 1858 . Pag. 58
P. rosenfeldi De Angeli & Garassino, 2006 . Pag. 58
Portunus Weber, 1795 . Pag. 58
P. arcuatus (A. Milne Edwards, 1860) . Pag. 58
P. convexus (Ristori, 1889) . Pag. 58
P. cfr. P. convexus (Ristori, 1889) . Pag. 59
94
ANTONIO DE ANGELI & ALESSANDRO GARASSINO
P. edwardsi E. Sismonda, 1846 . Pag. 59
P. hastatus (Linnaeus, 1767) . Pag. 59
P incertus (A. Milne Edwards, 1860) . Pag. 59
P. kochi (Bittner, 1893) . Pag. 59
P /arieti (A. Milne Edwards, 1860) . Pag. 59
P monspeliensis (A. Milne Edwards, 1860) . Pag. 59
P neogenicus Miiller, 1979 . Pag. 60
P. cf. P. radobojanus (Bittner, 1884) . Pag. 60
P cf. P stenaspis (Bittner, 1884) . Pag. 60
P. suessi (Bittner, 1875) . Pag. 60
P. tuberculatus Roux, 1 828 . Pag. 60
P vicentinus (A. Milne Edwards, 1860) . Pag. 61
Portunus (Achelous) De Haan, 1833 . Pag. 61
P. {Achelous) obtusus A. Milne Edwards, 1860 Pag. 61
Rakosia Miiller, 1984 . Pag. 61
R. grumìensis Beschin, De Angeli & Checchi,
2001 . Pag. 61
Scylla De Haan, 1833 . Pag. 61
Xanthoidea MacLeay, 1838 . Pag. 62
Carpiliidae Ortmann, 1893 . Pag. 62
Paìaeocarpilius A. Milne Edwards, 1 862 . Pag. 62
P. aquitanìcus A. Milne Edwards, 1862 . Pag. 62
P. macrochelus (Desmarest, 1 822) . Pag. 62
P. simplex Stoliczka, 1871 . Pag. 63
P. valrovinensis (De Gregorio, 1895) . Pag. 63
Domeciidae Ortmann, 1 893 . Pag. 63
Jonesius Sankarankutty, 1962 . Pag. 63
J. oligocenicus (Beschin, De Angeli & Checchi,
2001) . Pag. 64
Goneplacidae MacLeay, 1838 . Pag. 64
Branchioplax Rathbun, 1916 . Pag. 64
B. albertii De Angeli & Beschin, 2002 . Pag. 64
Chlinocephalus Ristori, 1886 . Pag. 64
C. demissifrons Ristori, 1886 . Pag. 64
Corallicarcinus Miiller & Collins, 1991 . Pag. 64
Gollincarcinus Beschin & De Angeli, 2004 . Pag. 64
G. levis Beschin & De Angeli, 2004 . Pag. 64
Goneplax Leach, 1 8 1 4 . Pag. 65
G. gulderi Bachmayer, 1953 . Pag. 65
G. rhomboides (Linnaeus, 1758) . Pag. 65
Lessinioplax Beschin & De Angeli, 2004 . Pag. 65
L. rugosa Beschin & De Angeli, 2004 . Pag. 66
L. simplex Beschin & De Angeli, 2004 . Pag. 66
Magyarcarcinus Schweitzer & Karasawa, 2004 Pag. 66
M. loczyanus (Lòrenthey, 1897) . Pag. 66
Maingrapsus Tessier, Beschin, Busulini & De
Angeli, 1999 . Pag. 66
M. quadratus Tessier, Beschin, Busulini & De
Angeli, 1999 . Pag. 66
Paracorallicarcinus Tessier, Beschin, Busulini
& De Angeli, 1999 . Pag. 66
P. arcanus Tessier, Beschin, Busulini & De
Angeli, 1999 . Pag. 66
Simonellia Vinassa de Regny, 1 897 . Pag. 66
S. quiricensis Vinassa de Regny, 1 897 . Pag. 67
Hexapodidae Miers, 1886 . Pag. 67
Stevea Manning & Holtuis, 1981 . Pag. 67
S. cesarli Beschin, Busulini, De Angeli & Tes¬
sier, 1994 . Pag. 67
Menippidae Ortmann, 1893 . Pag. 67
Eriphia Latreille, 1817 . Pag. 67
E. cocchii Ristori, 1 886 . Pag. 67
E. spinifrons (Herbst, 1782) . Pag. 67
Panopeidae Ortmann, 1893 . Pag. 68
Bittneria Schweitzer & Karasawa, 2004 . Pag. 68
B. attenuatus (Bittner, 1875) . Pag. 68
Carinocarcinus Lòrenthey, 1 898 . Pag. 68
C. zitteli Lòrenthey, 1898 . Pag. 68
Glyphithyreus Reuss, 1859 . Pag. 68
G. ellipticus (Bittner, 1875) . Pag. 68
Lophopanopeus Rathbun, 1898 . Pag. 68
Palaeograpsus Bittner, 1875 . Pag. 69
P injìatus Bittner, 1875 . Pag. 69
Panopeus H. Milne Edwards, 1834 . Pag. 69
P bolcensis De Zigno, 1915 . Pag. 69
P granulineatus Miiller & Collins, 1991 . Pag. 69
P vicentinus (Bittner, 1875) . Pag. 69
Pilumnidae Samouelle, 1819 . Pag. 70
Eohalimede Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 70
E. granosa Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 70
Eopilumnus Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 70
E. checchii Beschin, Busulini, De Angeli &
Tessier, 2002 . Pag. 70
Eumorphactaea Bittner, 1875 . Pag. 70
E. scissifrons Bittner, 1875 . Pag. 70
Galenopsis A. Milne Edwards, 1 865 . Pag. 70
G. crassifrons A. Milne Edwards, 1865 . Pag. 70
G. cfr. G. murchisoni A. Milne Edwards, 1865
. Pag. 70
G. ristorii Checchia-Rispoli, 1907 . Pag. 71
G. schopeni Checchia-Rispoli, 1905 . Pag. 71
G. similis Bittner, 1875 . Pag. 71
Lobogalenops is Miiller & Collins, 1991 . Pag. 71
L. quadrilobata (Lòrenthey, 1 897) . Pag. 7 1
Lobonotus A. Milne Edwards, 1864 . Pag. 72
L. cfr. L. orientalis Collins & Morris, 1978 . Pag. 72
Pilumnus Leach, 1815 . Pag. 72
P villosissimus (Rafinesque-Schmaltz, 1814) ... Pag. 72
Titanocarcinus A. Milne Edwards, 1863 . Pag. 72
T. aculeatus Busulini, Tessier & Visentin, 1984 Pag. 72
T. edwardsi (E. Sismonda, 1846) . Pag. 72
T. euglyphos Bittner, 1875 . Pag. 73
T. kochii Lòrenthey, 1897 . Pag. 73
T. raulinianus A. Milne Edwards, 1863 . Pag. 73
T. subovalis Ristori, 1 896 . Pag. 73
Xanthidae MacLeay, 1838 . Pag. 73
Monodaeus Guinot, 1967 . Pag. 73
M bortolottii Delle Cave, 1988 . Pag. 74
Paraxanthosia Miiller & Collins, 1991 . Pag. 74
CATALOG AND BIBLIOGRAPHY OF THE FOSSIL STOMATOPODA AND DECAPODA FROM ITALY
95
P. tuberculata Beschin, De Angeli, Checchi &
Zarantonello, 2005 . Pag.
Phlyctenodes A. Milne Edwards, 1862 . Pag.
P. dalpiazi Fabiani, 1911 . Pag.
?P. hantkeni Lòrenthey, 1898 . Pag.
IP. irregularis Ristori, 1896 . Pag.
P. krenneri Lòrenthey, 1 897 . Pag.
P. nicolisi Bittner, 1884 . Pag.
P. steinmanni Lòrenthey, 1901 . Pag.
Xantho Leach, 1 804 . Pag.
X. floridus Montagli, 1813 . Pag.
IX. lovisatoi (Lòrenthey, 1907) . . Pag.
X. moldavicus (Yanakevich, 1977) . Pag.
fZanthopsidae Via Boada, 1959 . Pag.
Harpactocarcinus A. Milne Edwards, 1 862 . Pag.
H. macrodactylus (H. Milne Edwards, 1850) .... Pag.
H. ovalis A. Milne Edwards, 1 862 . Pag.
H. punctulatus (Desmarest, 1 822) . Pag.
Harpactoxanthopsis Via Boada, 1959 . Pag.
H. quadrilobata (Desmarest, 1 822) . Pag.
H. cfr. H. quadrilobata (Desmarest, 1 822) . Pag.
H. souverbiei (A. Milne Edwards, 1 862) . Pag.
Neozanthopsis Schweitzer, 2003 . Pag.
N. bruckmanni (v. Meyer, 1 862) . Pag.
Potamoidea Ortmann, 1 896 . Pag.
Potamidae Ortmann, 1 896 . Pag.
Potamon Savigny, 1816 . Pag.
IP. castellinense (Szombathy, 1916) . Pag.
Thoracotremata Guinot, 1977 . Pag.
Pinnotheroidea De Haan, 1833 . Pag.
Pinnotheridae De Haan, 1833 . Pag. 79
Asthenognathus Stimpson, 1858 . Pag. 79
A. laverdensis De Angeli & Garassino, 2006 .... Pag. 79
Ocypodoidea Rafinesque-Schmaltz, 1815 . Pag. 79
Ocypodidae Rafinesque-Schmaltz, 1815 . Pag. 79
Archaeocypoda Secretan, 1975 . Pag. 79
A. veronensis Secretan, 1975 . Pag. 79
Palicidae Bouvier, 1898 . Pag. 79
Eopalicus Beschin, Busulini, De Angeli & Tes-
sier, 1996 . Pag. 79
E. imbricatus De Angeli & Beschin, 2000 . Pag. 79
E. semicarinatus De Angeli & Beschin, 2000 ... Pag. 79
E. squamosus Beschin, Busulini, De Angeli &
Tessier, 1996 . Pag. 79
Spinipalicus Beschin & De Angeli, 2003 . Pag. 80
S. italicus Beschin & De Angeli, 2003 . Pag. 80
Grapsoidea MacLeay, 1838 . Pag. 80
Grapsidae MacLeay, 1838 . Pag. 80
Daragrapsus Miiller & Collins, 1991 . Pag. 80
D. trispinosus Miiller & Collins, 1991 . Pag. 80
Daranyia Lòrenthey, 1901 . Pag. 80
D. fabianii Di Salvo, 1933 . Pag. 80
Grapsus Lamarck, 1801 . Pag. 80
Pseudodaranyia Tessier, Beschin, Busulini &
De Angeli, 1999 . Pag- 80
P carinata Tessier, Beschin, Busulini & De
Angeli, 1999 . Pag. 80
Acknowledgements . Pag. 81
Bibliography . Pag. 82
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Ili - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Al¬
pe Turati (Erba, Como). Generi Hildoceras, Phymatoceras, Paronice-
ras e Frechiella. Conclusioni generali, pp. 143-204, 2 figg., 6 tavv.
Volume XV II 1
I - PINNA G., 1969 - Revisione delle ammoniti figurate da Giuseppe Me¬
neghini nelle Tavv. 1-22 della « Monographie des fossiles du calcaire
rouge ammonitique» (1867-1881). pp. 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 tavv. n.t.
Ili - PETRUCCI F„ BORTOLAMI G. C. & DAL PIAZ G. V.. 1970 - Ri¬
cerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul
suo substrato cristallino, pp. 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., 1970 - Le Hildoceratidae del Lias medio delle regio¬
ni mediterranee - Loro successione e modificazioni nel tempo. Riflessi
biostratigrafici e sistematici, pp. 5-46, 2 tcibb. n.t.
II - PINNA G. & LEVI-SETTI F., 1971 - I Dactylioceratidae della Provin¬
cia Mediterranea (Cephalopoda Ammonoidea). pp. 47-136, 21 figg.,
12 tavv
III - PELOSIO G., 1973 - Le ammoniti del Trias medio di Asklepieion (Ar-
golide, Grecia) - I. Fauna del «calcare a Ptychites » (Anisico sup.). pp.
137-168, 3 figg., 9 tavv.
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,
37 figg., 6 carte tematiche.
Ili - CÈRETTI 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), pp. 5-34, 16 figg., 16 tavv.
II - POLUZZI A., 1975 - 1 Briozoi Cheilostomi del Pliocene della Val d’ Ar¬
da (Piacenza, Italia), pp. 35-78, 6 figg., 5 taw.
Ili - BRlAMBILLA G., 1976 - 1 Molluschi pliocenici di Villalvemia (Ales¬
sandria). 1. 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
III - 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„ 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, pp. 5-44, 4 figg., 16 tavv
II - POLUZZI A., 1982 - I Radiolari quaternari di un ambiente idrotermale
del Mar Tirreno, pp. 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 tavv.
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 1 830-1902. pp. 29-86, 24 figg., 12
tavv
Volume XXV
I - CALEGARI G., 1989 - Le incisioni rupestri di Taouardei (Gao, Mali).
Problematica generale e repertorio iconografico, pp. 1-14, 9 figg., 24
tavv
II - PINNA G. & NOSOTTI S., 1989 - Anatomia, morfologia funzionale
e paleoecologia del rettile placodonte Psephoderma alpinum Meyer,
1858. pp. 15-50, 20 figg., 9 tavv
III - CALDARA R., 1990 - Revisione Tassonomica delle specie paleartiche
del genere Tychius Germar (Coleoptera Curculionidae). pp. 51-218,
575 figg.
Volume XXVI
I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia, Placo-
dontia). pp. 1-21, 23 figg.
II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara preisto¬
rico: dati e interpretazioni. pp. 25-556, 647 figg.
HI - ANDR1 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 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., 1999 - L’arte rupestre dell’Eritrea. Repertorio ragionato
ed esegesi iconografica, pp. 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, pp. 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). pp. 1-31, 21 figs., 9 pls.
Volume XXXI
I - NOSOTTI S. & RIEPPELO., 2002 - The braincase of Placodus Agassiz,
1833 (Reptilia, PIacodontia).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 tavv.
Ili - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphargis dalsassoi n. gen.
n. sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano
(Lombardy, N Italy). pp. 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). 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. 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.
II - FUCHS D., 2006 - Morphology, taxonomy and diversity of vampyropod
Coleoids (Cephalopoda) from thè Upper Cretaceous of Lebanon. pp.
1-28, 9 figs., 9 pls.
Ili - CALDWELL M. W., 2006 - A new species of Pontosaurus (Squamata,
Pythonomorpha) from thè Upper Cretaceous of Lebanon and a phylo-
genetic analysis of Pythonomorpha. pp. 1-42, 18 figs., 1 pi.
Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali,
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Pubblicazione disponibile al cambio
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looà
Volume XXXV - Fascicolo li
MCZ
UBRARV
,1W 1 3 2007
della Società Italiana
di Scienze Naturali
e del Museo Civico
di Storia Naturale di Milano
3rd SYMPOSIUM ON MESOZOIC AND CENOZOIC
DECAPOD CRUSTACEANS
Museo di Storia Naturale di Milano
May 23-25, 2007
Edited by
ALESSANDRO GARASSINO
RODNEY M. FELDMANN
GIORGIO TERUZZI
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MILANO APRILE 2007
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 jacobita (Com.). 9 pp., 1 tav.
II - MAGNI-GRIFFI F., 1865 - Di una specie d 'Hippolais nuova per l’Ita¬
lia. 6 pp., 1 tav.
Ili - GASTALDI B., 1 865 - 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 /aw.
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., 1 865 - Intorno al genere Aeolosoma. 18 pp., 2 taw.
X - CORNALIA E., 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., 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 taw.
Ili - MOLON F., 1 867 - 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., 1 866 - 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.
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 - DÙRER 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 Vìpera Redii. 16 pp., 1 tav.
II - GARO VAGLIO S., 1867 - Thelopsis, Belonia, Weitenwebera et Limbo¬
ria, quatuor Lichenum Angiocarporum genera recognita iconibusque
illustrata. 12 pp., 2 taw.
III - 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 - 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 nummulitico del-
l’ Alpi venete. Parte 1 1 . 32 pp. 8 taw.
II - GARO VAGLIO S., 1868 - Octona Lichenum genera vel adhuc con¬
troversa, vel 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 - MARTORELLI G., 1898 - Le forme e le simmetrie delle macchie nel
piumaggio. Memoria ornitologica. 112 pp., 63 figg., 1 taw.
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 petrografia e
geologici. Parte I. pp. 1-46, 5 figg., 3 taw.
II - REPOSSI E., 1916 (1917) - La bassa Valle della Mera. Studi petrogra¬
fia e geologici. Parte II .pp. 47-186, 5 figg. 9 taw.
Ili - AIR AGHI 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 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, 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.
Ili - SCORTECCI G„ 1941-1 recettori degli Agamidi. pp. 209-326, 80 figg.
Volume XI
I - GUIGLIA D., 1 944 - 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.
1 - 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.
Ili - VIALLI V., 1959 - Ammoniti sinemuriane del Monte Albenza (Berga¬
mo), pp. 141-188, 2 figg., 5 taw.
Volume XIII
I - VENZO S., 1961- Rilevamento geologico dell’anfiteatro morenico del
Garda. Parte II. Tratto orientale Garda-Adige e anfiteatro atesino di
Rivoli veronese, pp. 1-64, 25 figg., 9 taw, 1 carta.
II - PINNA G., 1963 - Ammoniti del Lias superiore (Toarciano) dell’Alpe
Turati (Erba, Como). Generi Mercaticeras, Pseudomercaticeras e Bro-
dieia. 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., 8 taw.
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 - 11 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, 27 figg. 9 taw.
II - DIENI I. e MASSARI F., 1 966 - Il Neogene e il Quaternario dei dintorni
di Orosei (Sardegna), pp. 89-142, 8 figg., 7 taw.
Ili - BARBIERI F., 1ACCARINO S., BARBIERI F. & PETRUCCI F„ 1967
- Il Pliocene del Subappennino Piacentino-Parmense-Reggiano. pp.
143-188, 20 figg., 3 taw.
Volume XVI
I - CARETTO P. G., 1967 - Studio morfologico con l’ausilio del metodo
statistico e nuova classificazione dei Gasteropodi pliocenici attribuibili
al Mure \ 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 - 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,
llammatoceratidae (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-
FAnisico superiore di Lenna in Val Brembana (Bergamo), pp. 71-142,
5 figg., Il taw.
3rd Symposium
on Mesozoic and Cenozoic Decapod Crustaceans
Museo di Storia Naturale di Milano
May 23-25, 2007
Edited by
Alessandro Garassino
Rodney M. Feldmann
Giorgio Teruzzi
MCZ
library
JUN 1 3 2007
HARVARD
UNIVERSITY
1857 - 2007
150 ANNI
DI NATURA
E SCIENZA
SOCIETÀ ITALIANA
DI SCIENZE NATURALI
Volume XXXV - Fascicolo II
Aprile 2007
Memorie della Società Italiana di Scienze Naturali
e del Museo Civico di Storia Naturale di Milano
INDEX
Foreword by Feldmann R. M . Pag. 3
Ahyong S. T. & Garassino A. - A new stomatopod from
thè Upper Cretaceous (Cenomanian) ofLebanon . Pag. 5
Artal P. & Gilles A. - New Miocene crabs from Pignan
(southeast France) . Pag. 8
Beschin C., Busulinì A. & Tessier G. - First report of a
new Eocene crustacean fauna from thè Veronese Lessini
(N Italy) . Pag. 12
Bishop G. A. - Mechanism for phosphatization of Dakoti-
cancer assemblages in thè Late Cretaceous western interior
Seaway, USA . Pag. 15
Breton G. & Collins J. S. H. - Decapod fauna from thè
Cenomanian stratotype . Pag. 17
De Angeli A. & Beschin C. - Tertiary stomatopods from
Italy . Pag. 21
De Angeli A. & Garassino A. - Decapod crustaceans from
thè Mesozoic and Cenozoic of Friuli-Venezia Giulia (NE
Italy) . Pag. 24
Feldmann R. M., Green R. & Schweitzer C. E. -
An unusual Albian (Early Cretaceous) crab from Montana:
thè earliest eubrachyuran? . Pag. 28
Feldmann R. M. & Schweitzer C. E. - A compilation of
Decapoda collections in Museums of thè world . Pag. 30
Fraaije R. H. B., Van Bakel B. W. M. & Jagt J. W. M. - A
new species of Gonìocypoda and thè first record of Glyphi-
thyreus wetherelli (Bell, 1858) (Decapoda, Brachyura)
from thè Eocene of Nieuwvliet-Bad, The Netherlands . Pag. 37
Garassino A. & De Angeli A. - Report of Ranilia constricta
(A. Milne Edwards, 1880) (Brachyura, Raninidae) from
thè Tyrrhenian (upper Pleistocene) of Bovetto (Calabria, S
Italy) . Pag. 43
Garassino A., De Angeli A. & Pasini G. - New decapod
assemblage from thè Upper Cretaceous (Cenomanian-Turo-
nian) of Gara Sbaa, southeastem Morocco . Pag. 45
Giannetti A., Monaco P., Caracuel J. E., Soria J. M. &
Yébenes A. - Functional morphology and ethology of deca¬
pod crustaceans gathered by Thalassinoides branched bur-
rows in Mesozoic shallow water environments . Pag. 48
Guinot D., De Angeli A. & Garassino A. - Discovery
of thè oldest eubrachyuran crab from thè Middle Jurassic
(Bathonian) of Normandy (France) . Pag. 53
Hernàndez-Monzón Ó., Vega F. J. & Coutino M. A. - A
review of Lophoranina cristaspina from thè Middle Eocene
of Chiapas, Mexico and evolutionary implications . Pag. 56
Hyzny M. - Paleogene crab fauna of Borové Formation
(localities Durkovec and Hlinisko), western Carpathians,
Slovakia . Pag. 59
Karasawa H. & Rato H. - New prosopid crabs (Crustacea,
Decapoda, Brachyura) from thè Upper Jurassic Torinosu
Group, Shikoku, Japan . Pag. 62
Krobicki M., Miiller P. & Zaton M. - Middle and Upper
Jurassic European prosopid crabs, phylogeny and palaeoen-
vironments . Pag. 66
Larghi C. & Tintori A. - First report of a decapod from
thè Meride Limestone: new data from one of thè best
Ladinian (Middle Triassic) taphonomic Windows of a tran-
sitional environment . Pag. 68
Marangon S. & De Angeli A. - Preservation of some speci-
mens of Portunus monspeliensis (A. Milne Edwards, 1860)
from thè Miocene of Sardinia (Italy) . Pag. 70
Marangon S. & De Angeli A. - New decapod assemblage
from thè lower Oligocene (Rupelian) of Bacino Ligure Pie¬
montese (N W Italy) . Pag. 73
Martins-Neto R. G. & Dias Junior S. C. - The Brazilian
paleodecapod fauna: state of thè knowledge . Pag. 76
Monaco P., Caracuel J. E., Giannetti A., Soria J. M. &
Yébenes A. - Thalassinoides and Ophiomorpha as cross-
facies trace fossils of crustaceans from shallow-to-deep-
water environments: Mesozoic and Tertiary examples from
Italy and Spain . Pag. 79
Neto de Carvalho C. & Viegas P. A. E. J. - Microfacies
analysis of Thalassinoides infilling: causes of collective burial
among Mecochirus populations . Pag. 83
Robins C. M., Feldmann R. M. & Schweitzer C. E. - Prim¬
itive brachyurans and galatheids from Emstbrunn, Austria:
an evaluation of thè Friedrich Bachmayer Collection . Pag. 85
Schweigert G. - Preservation of decapod crustaceans in
thè Upper Jurassic lithographic limestones of southern Ger-
many . Pag. 87
Schweitzer C. E., Feldmann R. M., Shirk A. M. &
Lazàr I. - Differential diversity in Jurassic sponge-algal
and coralline communities, Romania . Pag. 91
Teruzzi G. & Garassino A. - Coleia Broderip, 1835 (Cru¬
stacea, Decapoda, Coleiidae) from thè Mesozoic of Italy: an
update . Pag. 95
Vega F. J., Àlvarez F. & Carbot-Chanona G. - Albian
penaeoidea (Decapoda, Dendrobranchiata) from Chiapas,
southern Mexico . Pag. 97
Verhoff J. R., Miiller P. M., Feldmann R. M. «Se Sch¬
weitzer C. E. - A novel Paleocene decapod fauna from thè
Kambuhel Formation . Pag. 101
Waugh D. A., Feldmann R. M., Hull A., Hein K. & Sch¬
weitzer C. E. - Scaling and classification of cuticular pits in
raninids . Pag. 103
© 2007 Società Italiana di Scienze Naturali
Museo Civico di Storia Naturale di Milano
Corso Venezia ,55 -20121 Milano
In copertina: Rosenfeldia triasica Garassino, Teruzzi & Dalla Vecchia, 1996 (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 - Aprile 2007
ISSN 0376-2726
Interest in thè paleontological history of thè deca-
pod Crustacea has exploded within thè past three dec-
ades. Perhaps stimulated by thè publication, in 1969, of
Glaessner’s Decapoda in thè Treatise on Invertebrate
Paleontology, thè number of people devoted to thè study
of decapod fossils and thè number of published papers on
thè subject mushroomed. By thè beginning of this Cen-
tury, there had been so many advances in research that
thè Treatise was beginning to show its age, and plans had
been laid to produce a revision of that landmark reference.
Recognizing that thè field had become so vast and that
thè range of taxa was so great, it was decided to invite
experts in different aspects of thè study of decapods,
from all over thè world, to participate in thè revision.
To launch thè effort, thè lst Workshop on Mesozoic and
Tertiary Decapod Crustaceans was organized and held
in Montecchio Maggiore, Vicenza, Italy. The meeting
was attended by over 30 people from all over thè world,
papers were delivered on a range of subjects from pale-
oecology, ichnology, systematic paleontology, and theory
of classification. Those in attendance included nearly all
thè most active workers on decapod fossil living at that
time. The meeting was wonderful, thè response was out-
standing, and thè enthusiasm for continuing thè work on
thè group was contagious! At thè dose of thè meeting, it
was decided to hold another meeting on thè subject, and
Boxtei, Maastricht, The Netherlands, was selected as thè
meeting site. That session, held in September, 2003, was
attended by essentially thè same group of people with thè
addition of some new students. One of thè conclusions
reached at that meeting was that thè Treatise revision was
stili in progress, but that thè volume of research published
since thè first meeting was so great that much more work
needed to be done. I believe it is fair to say that thè meet¬
ing in Boxtei was as enthusiastically received as was thè
first and, again, we went away with thè pian to continue
thè tradition of discussing thè decapods.
Now, we are at thè 3rd Symposium and thè excitement
about thè decapods continues to grow. As one metric, a
quick examination of thè papers retrievable on thè Georef
search engine of thè American Geological Institute indi-
cates that over 120 papers on fossil decapods have been
published since thè last meeting in 2003, and that number
does not include thè majority of papers published within
2006! Examination of some of thè titles reveals that these
have not been trivial papers - entire families and super-
families have been revised, large compilations of data on
paleobiogeography and been published, and extremely
useful catalogues of regional collections and classic
faunas are now available. Thus, thè work on thè Deca¬
poda is proceeding at an unprecedented rate. The attend¬
ance at this meeting, based upon pre-registration figures,
promises to be as large as thè previous meetings, thè geo-
graphic spread of attendees is as great, and thè range of
papers published in this volume of extended abstracts is
broad and thè topics continue to be exciting.
Finally, we all owe a debt of gratitude to Alessandro
Garassino for his tireless efforts in making this meeting
possible as well as to those who organized thè previous
two meetings. I am confident that thè 3rd Symposium on
Mesozoic and Cenozoic Decapod Crustaceans will be a
success and that it will be another in a very long series
of meetings. Enjoy thè abstracts, enjoy thè presentations,
and perhaps most important, enjoy thè company of your
colleagues and friends in this wonderful setting.
Rodney M. Feldmann
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
-
—fri ■ ~ir~
Shane T. Ahyong & Alessandro Garassino
A new stomatopod from thè Upper Cretaceous (Cenomanian)
of Lebanon
To date, two species of stomatopod are known from
thè Upper Cretaceous (Cenomanian) of Lebanon: Pseu¬
dosculda laevis (Schiuder, 1 872) (Pseudosculdidae Dames,
1886) and Sculda syriaca Dames, 1886 (Sculdidae Dames,
1 886), studied by Schiùder ( 1 872, 1 874), Woodward ( 1 879),
Dames (1886), Roger (1946), and Teruzzi (1983). Sculdi¬
dae and Pseudosculdidae, with a Jurassic-Cretaceous and
Upper Cretaceous range, respectively, are believed to rep-
resent stem-lineage unipeltatans (see Hof, 1998; Ahyong &
Harling, 2000). An unusual fossil stomatopod was discov-
ered in thè sedimentary layers of Haqel quarry (northeast-
em Lebanon), of lower-middle Cenomanian age (Upper
Cretaceous) (Hiickel 1970, 1974a, 1974b). Haqel, located
about 1 2 km from thè Mediterranean coast and 45 km from
Beirut, is one of thè richest fossil sites in Lebanon, yield-
ing numerous vertebrate and invertebrate fossils, including
Crustacea (e.g. Dames, 1886; Glaessner, 1945; Roger,
1946; Garassino, 1994, 2001; Larghi, 2004; Garassino &
Schweigert, 2006).
Ahyong et al. (2007) described Archaeosculda phoe-
nicia, thè second known genus and species of Pseudo¬
sculdidae. It has a subovate telson and is slightly broader
than long, with a distinct median carina and four pairs
of primary marginai teeth of which thè submedians have
movable apices, and a broad, subovate uropodal endopod.
This species is unique in thè Pseudosculdidae by exhib-
iting a broad telson with three pairs of prominent, fixed
primary teeth (in addition to thè moveable submedian
teeth) and a broad, ovate, uropodal endopod. In contrast,
thè only other recognised pseudosculdid, Pseudosculda
laevis , bears an elongate, triangular telson with small
primary teeth that do not project much beyond thè generai
telson outline, and a slender, elongate, uropodal endopod.
Moreover, at 75 mm total length, A. phoenicia apparently
reaches a larger size than P. laevis, known to reach only
55 mm total length (Holthuis & Manning, 1969). The
dissimilarities between thè telson of Archaeosculda and
Pseudosculda are marked, but thè four pairs of primary
teeth can probably be homologized between thè two
genera. Homologizing thè telson teeth of pseudosculdids
and those of crown-group unipeltatans is more compli-
cated, and requires further research.
Feldmann et al. (1999) reported a fragmentary fossil
stomatopod from Cretaceous of Colombia that was tenta-
tively assigned to Sculda on thè basis of telson morphol-
ogy whereby thè length and width are approximately
equal and thè posterior margin is lined with long, move¬
able spines, a distinctive feature of Sculdidae. Schram
& Miiller (2004), however, reassigned thè Colombian
specimen to Pseudosculda on thè basis of thè raptorial
claw. The dactyli of thè raptorial claws of thè Colombian
specimen resemble those of pseudosculdids, but it is
noteworthy that thè raptorial claws of Sculda are pres-
ently unknown. The Colombian specimen is certainly not
Pseudosculda, and it is doubtful that it belongs to Pseu¬
dosculdidae either.
Cladistic analyses of thè Stomatopoda have recognised
Sculdidae and Pseudosculdidae as stem-lineage unipelta¬
tans, with thè pseudosculdids as sister to crown-group
Unipeltata (see Hof, 1998; Ahyong & Harling, 2000).
The chief character excluding sculdids and pseudosculd¬
ids from thè crown-group is thè plesiomorphic uropodal
exopod segmentation, being unisegmental rather than
bi-segmental. Sculdidae, as presently composed, can be
diagnosed by several apomorphies such as thè broad,
dorsoventrally depressed body, thè absence of a median
carina on thè telson, and most importantly, thè posterior
telson margin lined with moveable spines. Unfortunately,
thè raptorial claws of sculdids, being unknown, cannot be
compared to those of pseudosculdids. Pseudosculdidae
is distinguished from Sculdidae by thè narrower, more
subcylindrical body form, and thè median carina on thè
telson. Only thè submedian primary telson teeth are artic-
ulated. Hof (1998) and Schram & Miiller (2004) regarded
thè well-sclerotized scythe-like dactylus of thè raptorial
claw as a defining feature of Pseudosculdidae. The rapto¬
rial claws of members of thè crown-group Hemisquillidae
Manning, 1980 (superfamily Gonodactyloidea), however,
are structurally similar to pseudosculdids, albeit with
proportionally shorter dactyli and propodi. The differ-
ences between thè raptorial claws of hemisquillids and
pseudosquillids are essentially morphometric, so care
should be taken to avoid placing undue emphasis on thè
raptorial claws in defining and assigning taxa to Pseu¬
dosculdidae. Note also that Hemisquilla Hansen, 1895,
is basai in thè Gonodactyloidea (Ahyong & Harling,
2000), so its raptorial claw morphology perhaps reflects
thè stem-lineage condition. Other recognized characters
of Pseudosculdidae are not unique - thè subcylindrical
body is a plesiomorphy inherited from archaeostomat-
opodeans and retained by thè modem gonodactyloids
and parasquilloids; and thè telson with a distinct median
carina and moveable submedian telson teeth is retained by
almost all modem unipeltatans. Consequently, no robust
synapomorphies are presently known that unite Pseu¬
dosculda and Archaeosculda raising thè possibility that
Pseudosculdidae is paraphyletic. To be sure, thè anterior
cephalic appendages of Pseudosculda and Archaeosculda
are incompletely known and more complete fossils may
reveal synapomorphies. Nevertheless, thè discovery
of Archaeosculda phoenicia is significane not only for
expanding thè rare Mesozoic stomatopod fossil record,
but also in increasing knowledge of thè morphospace
occupied by stem-lineage unipeltatans.
6
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
At present, sculdids are known reliably from thè
Upper Jurassic to Middle Cretaceous and thè pseudo-
sculdids from thè Upper Cretaceous. Pinna (1985) listed
two ill-preserved stomatopod fossils from thè Lower
Jurassic (Sinemurian) of Osteno in northern Italy, rep-
resenting thè oldest known unipeltatans. Hof (1998)
indicated that thè Osteno specimens represent an unde-
scribed Pseudosculda- like species. If thè Osteno speci¬
mens are pseudosculdids, and if current phylogenetic
hypotheses are correct, then both thè Pseudosculdidae
and Sculdidae are considerably older than presently
recognized.
Fig. 1 — A) A. phoenicia Ahyong, Garassino & Gironi, 2007, dorso-lateral view. B) A. phoenicia Ahyong, Garassino & Gironi, 2007
tail fan. C) Recostruction of thè tail fan.
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
7
References
Ahyong S. T. & Harling C., 2000 - The phylogeny of thè
stomatopod Crustacea. Austr aliati Journal ofZoology,
48 (6): 607-642.
Ahyong S. T., Garassino A. & Gironi B., 2007 - Archaeo-
sculda phoenicia n. gen., n. sp. (Crustacea, Stomat-
opoda, Pseudosculdidae) from thè Upper Cretaceous
(Cenomanian) of Lebanon. Atti della Società italiana
di Scienze naturali e del Museo civico di Storia natu¬
rale in Milano , Milano, 148 (1): 3-15.
Dames W., 1 886 - Ueber einige Crustaceen aus den Kreid-
ablagerunden des Libanon. Zeitschrift der Deutschen
Geologischen Gesellschaft, Berlin, 38: 551-575.
Feldmann R. M., Villamil T. & Kauffman E. G., 1999 -
Decapod and stomatopod crustaceans from mass mor-
tality Lagerstatten: Turonian (Cretaceous) of Colom¬
bia. Journal ofPaleontology, Lawrence, 73: 91-101.
Garassino A., 1994 - The macruran decapod crustaceans
of thè Upper Cretaceous of Lebanon. Paleontologia
Lombarda , Nuova serie, Milano, 3: 3-27.
Garassino A., 2001 - New decapod crustaceans from thè
Cenomanian (Upper Cretaceous) of Lebanon. Atti
della Società italiana di Scienze naturali e del Museo
civico di Storia naturale in Milano, Milano, 141 (2):
237-250.
Garassino A. & Schweigert G., 2006 - Cretasergestes
sahelalmaensis n. gen., n. sp. (Crustacea, Decapoda,
Sergestidae) and Cancrinos libanensis n. sp. (Crusta¬
cea, Decapoda, Palinuridae) from thè Late Cretaceous
(Cenomanian) of Lebanon. Atti della Società italiana
di Scienze naturali e del Museo civico di Storia natu¬
rale in Milano, Milano, 147 (1): 69-78.
Glaessner M. F., 1945 - Cretaceous Crustacea from
Mount Lebanon, Syria. Annals and Magazine of Natu¬
rai History Museum, London, 12 (11): 694-707.
Hof C. H. J., 1998 - Fossil stomatopods (Crustacea:
Malacostraca) and their phylogenetic impact. Journal
of Naturai History, 32: 1567-1576.
Holthuis L. B. & Manning R. B., 1969 - Stomatopoda.
In: Treatise on Invertebrate Paleontology. Arthropoda.
Moore R. C. (ed.). Geologica l Society of America and
University of Kansas, Lawrence, 4 (2): R535-552.
Hiickel U., 1970 - Die Fischschiefer von Hakel und
Hjoula in der Oberkreide des Libanon. Neues Jahr-
buch fùr Geologie und Palàontologie, Abhandlungen,
Stuttgart, 135 (2): 113-149.
Hiickel U., 1974a - Vergleich des Mineralbestandes der
Plattenkalke Solnhofens und des Libanon mit anderen
Kalken. Neues Jahrbuch fùr Geologie und Palàonto¬
logie, Abhandlungen, Stuttgart, 145 (2): 153-182.
Hiickel U., 1974b - Geochemischer Vergleich der Plat¬
tenkalke Solnhofens und des Libanon mit anderen
Kalken. Neues Jahrbuch fùr Geologie und Palàonto¬
logie, Abhandlungen, Stuttgart, 145 (3): 279-305.
Larghi C., 2004 - Brachyuran decapod crustaceans from
thè Cenomanian (Upper Cretaceous) of Lebanon.
Journal ofPaleontology, Lawrence, 78 (3): 528-541.
Pinna G., 1985 - Exceptional preservation in thè Jurassic
of Osteno. Philosophical Transactions of thè Royal
Society of London, London, B 3 1 1 : 171-180.
Roger J., 1946 - Les invertébrés des couches a poissons
du Crétacé supérieur du Liban. Mémoires de la Société
Géologique de France, Paris, 23: 1-92.
Schliiter C., 1872 - Ueber einen fossilen Stomatopoden
von Libanon. Sitzungberichte Naturistorischen Verein
der Preussischen Rheinlande und Westfalens, Bonn,
29: 194-195.
Schliiter C., 1874 - Ueber einige jurassische Crustaceen-
Typen in der oberen Kreide. 1. Fossile Krebse des
Libanon. Sitzungberichte Naturistorischen Verein der
Preussischen Rheinlande und Westfalens, Bonn, 31:
41-55.
Schram F. R. & Miiller H.-G., 2004 - Catalog and bibli-
ography of thè Fossil and Recent Stomatopoda. Back-
huys publishers, Leiden, The Netherlands.
Teruzzi G., 1983 - Un nuovo esemplare di Palaeosculda
laevis (Schliiter, 1872) del Cenomaniano di Hakel nel
Libano. Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale in Milano, Milano,
124(1-2): 117-122.
Woodward H., 1879 - Contributions to thè knoweledge of
fossil Crustacea. Quarterly Journal of thè Geological
Society, London, 35: 549-556.
Shane T. Ahyong - National Institute of Water and Atmospheric Research, Private Bag 14901, Kilbimie, Wellington, New Zealand.
e-mail: s.ahyong@niwa.co.nz
Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: agarassino@libero.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Pedro Artal & Alonso Gilles
New Miocene crabs from Pignan (southeast France)
Marine sediments from thè Chàteau Saint-Martin
quarry, stili active, situateci dose to thè village of Pignan,
and not far from Montpellier (Herault, southeast France),
have yielded a very rich crustacean fauna during thè
last fi ve years. The outcrops are frequently removed by
mining, and are nearly at thè point of disappearing. We
have recorded a decapod association characteristic for
thè Mediterranean area. Miocene strata around Montpel¬
lier has been studied from thè XIX century (see Roman,
1897), and were first mentioned in terms of remains of
fishes, mainly shark teeth, and sporadic crabs of large
size. The upper sediments from thè area, thè subject of
this note, are about 20 m thick, and consist of bioclastic
carbonates with embedded pockets of laminated clayey
silts containing some fossil plants. The silt pockets are
3 m thick and about 10 m long. The underlaying beds,
composed of sands, conglomerates of small size, and
marls, yielded an abundant and diverse fìsh fauna. This
fauna completely corresponds to thè fauna of selachians
described by Capetta (1970), which were collected from
sediments of Helvetian and Burdigalian age in thè Her¬
ault. The authors of thè geological map of Montpellier
mention of thè Pignan quarry and thè bioclastic layers
as being upper Burdigalian in age. The shell calcaren-
ites dominated by moulds of gastropods, bivalves, some
bryozoans, and worms provided very few crabs, except
for some small calappids and leucosids. Most crabs were
collected in thè clayey and sandy silt pockets, being
better preserved according to thè grain size. In thè basai
beds rich in fish remains Necronectes sp. and Portu¬
nus monspeliensis (A. Milne Edwards, 1860) of major
dimensions occur.
The decapod assemblage contained in thè clay mainly
consists of isolated carapaces of average to small size,
from a few milimeters to five centimeters. Corpses are
extremely rare, except for some portunids and goneplac-
ids. All thè assemblage has been tentatively assigned to 13
families and 16 genera. The most frequent, in number of
specimens, are portunids, calappids, leucosids, dorippids,
and majids. Some other genera are only represented by
a unique fossil, very few specimens, or badly preserved
remains.
Portunids, thè most common and diverse decapods,
tentatively include thè genera Portunus Weber, 1795,
Liocarcimis Stimpson, 1871, and Necronectes A. Milne
Edwards, 1881. They tend to be small size. The family
Calappidae H. Milne Edwards, 1837, appears to be very
abundant, with very well preserved small carapaces of
thè genus Calappa Weber, 1795. The family Dorippidae
MacLeay, 1838, is well represented in number of speci¬
mens of thè genus Ethusa Roux, 1830, rather Constant,
and probably thè genus Medorippe Manning & Holthius,
1981. Leucosiids are also common and rather well pre¬
served, thè family Leucosiidae Samouelle, 1819, is doc-
umented by thè genus Ebalia Leach, 1817; distinctive
isolated female abdomina also occur. The family Maji-
dae Samouelle, 1819, is represented by only one genus,
frequent in number despite thè shell not being well
preserved. The goneplacids, very common in European
Miocene strata are also present; in this case thè family
Goneplacidae Mac Leay, 1838, can be documented
with confidence through thè genus Goneplax Leach,
1814. Xanthoids belonging to thè families Panopeidae
Ortmann, 1893, and Menippidae Ortmann, 1893, are
confirmed by well preserved material of thè genera Pan-
opeus H. Milne Edwards, 1834, and Eriphia Latreille,
1817. Dromiids and cancroids are rare but certain; thè
genus Atelecyclus Leach, 1814, is confirmed by a unique
specimen of unusually small size. Some homolids and
possible pirimelids also seem to be present, but better
specimens are needed for confìrmation. The absence of
isolated claws, and thè absence of anomurans is remark-
able. Fingers and more or less complete palms are hard
parts considered to be easily preserved. Only a large
finger of a possible pagurid has been recorded to date.
The decapod association, thè assemblage of molluscs,
and thè terrigenous sediments containing fossil leaves,
suggest a littoral environment. The fossil plants are sim-
ilar to thè genus Avicennia, which suggests thè possible
proximity of a mangrove.
Miocene crab faunas are not well represented in
thè south of Europe, except for thè rich Badenian of
Hungary (Miiller, 1984) and less diverse, but also
abundant, from Catalonia (Miiller, 1993; Solé & Via,
1988). Miocene decapods from Italy consist of small
assemblages of few genera (De Angeli & Marangon,
1992; De Angeli & Garassino, 2006). The new French
decapod association cannot be completely related to thè
former assemblages, probably due to thè provenance
from different environments, but it does share some
genera with thè faunas from Italy, Spain, and Hungary.
In Catalonia, thè nearest area with similar Miocene
layers, Portunus monspeliensis is also common, but
occurs occasionally associated with other decapods or
in less diverse assemblages. The frequent Macrophthal-
mus aquensis A. Milne Edwards & Brocchi, 1879 (see
Miiller, 1993) is not present in Pignan. The entire new
assemblage from thè southeast of France, at thè level
of genera, seems to be confirmed as strongly related
to thè recent Mediterranean fauna (Zariquiey, 1968).
Further studies and possible new material improving
our understanding of thè poorly preserved genera will
permit increasing our knowledge of thè Mediterranean-
Thetyan realm.
3R1) SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
9
Fig. 1 - A) Necronectes cf. schajferi. B) Liocarcinus sp. C) Portunus monspeliensis, ventral view. D) Necronectes sp. E) Eriphia sp.
F) Goneplax sp.
10
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Fig. 2 - A) Calappa sp. B) Ebalia sp., dorsal view. C) Ebalia sp., abdomen. D) Panopeus sp. E) Majidae sp. F) Ethusa sp.
3RI> SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
11
Fig. 3 - A) Dromiidae sp. B) Atelecyclus sp. C) Dorippidae sp. D) Portunidae sp.
References
Capetta H., 1970 - Les sélaciens du Miocène de la région
de Montpellllier. Palaeovertebrata mém. ext., Mont¬
pellier: 1-139.
De Angeli A. & Garassino A., 2006 - Catalog and biblio-
graphy of thè fossil Stomatopoda and Decapoda from
Italy. Memorie della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale di Milano,
Milano, 35 (1): 1-95.
De Angeli A. & Marangon S., 1992 - Necronectes schaf-
feri Glaessner, nel Miocene della Sardegna (Italia).
Lavori - Società Veneziana di Scienze Naturali, Ve¬
nezia, 17: 175-182.
Milne Edwards A., 1881 - Notes sur quelques crustacés
fossiles des environs de Biarritz. Annales des Sciences
Geologiques, 11 (2): 1-8.
Mtiller P., 1984 - Decapod Crustacea of thè Badenian.
Geologica Hungarica, Serie Palaeontologica, 42:
1-121.
Miiller R, 1993 - Neogene decapod crustaceans from
Catalonia. Scripta Musei Geologici Seminarii Barci-
nonensis, Barcelona, 225: 1-39.
Roman F., 1897 - Recherches stratigraphiques et paléon-
tologiques dans le Bas-Languedoc. Annales Univer¬
sità Lyon, Ly on: 1-345.
Solò J & Via L., 1988 - Crustacis Dècapodes fòssils dels
Pai'sos Catalans (Recopilació i actualització de dades
de 1855 a 1988). Batalleria, Barcelona, 2: 23-42.
Zariquiey Alvarez R., 1968 - Crustàceos Decàpodos
Ibéricos. Investigaciones Pesqueras, Barcelona, 32:
1-510.
Pedro Artal - Museu Geologie del Seminari Conciliar de Barcelona, Diputación 23 1 , 08007 Barcelona, Spain.
e-mail: partal@optimus.es
Alonso Gilles - Chemin du mas de Daulet 3, 34570 Pignan, France.
e-mail: saintseiyar64@hotmail.com
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Claudio Beschin, Alessandra Busulini & Giuliano Tessier
First report of a new Eocene crustacean fauna
from thè Veronese Lessini (N Italy)
Parona is a little village located near thè western bound-
ary of thè Verona commune, in thè last part of thè Negrar
Valley (Fig. 1). Here an interesting fossil crustacean fauna
has been found in thè Priabonian levels that now are no
more accessible because of intense urbanization.
From a morphological point of view, this area is
characterized by a series of low hilly digitations directed
N-S stretching out toward thè Po River and Veneto plain.
The plain is here made of thè detritai materials carried by
Adige River that flows in meanders near thè slopes of thè
same low hills.
The Parona stratigraphic series is Eocene in age; it
is comprised of bedded carbonate deposits (Nummulite
Limestones of middle Eocene), on which is overlain grey-
greenish and/or yellowish marls without clear stratifica-
tion and full of discocyclinids (Priabona Marls of upper
Eocene). On thè ground, thè limits of these two units are
very apparent above all by thè presence on thè upper part
of thè series of thè Priabonian clayey lithotypes.
From a palaeontological point of view in thè studied
Priabonian formation, larger foraminifera and bivalves
(particularly ostreids), but also coralline algae, bryozoans,
gastropods, echinids, and crustaceans have been found.
The fossil association indicates a nerithic shallow marine
environment probably not far from thè coast and affected
by significant terrigenous supplies (De Zanche et al. , 1 977).
The analysis of thè fossil crustaceans found here and
now deposited in thè Museo Civico “G. Zannato” at Mon-
tecchio Maggiore (Vicenza), allowed identification of about
ten species of Brachyura; thè remains consist of carapaces
and some propoda of chelipeds. Their preservation is not
good; thè originai shell is often altered and appears dusty.
Even when thè number of specimens is moderate, thè
fauna is differentiated: thè following genera have been
identified within seven families: Dromiidae de Haan,
1833; Noetlingia Beurlen, 1928: Dynomenidae Ortmann,
1892; Kromtìtis Miiller, 1984: Raninidae de Haan, 1841;
Lophoranina Fabiani, 1910: Calappidae de Haan, 1833;
Calappilia A. Milne Edwards, 1873: Parthenopidae
MacLeay, 1838; “ Phrynolambrus ” Bittner, 1893: Pilum-
nidae Samouellel819; Eohalimede Blow & Manning,
1996; Lobonotus A. Milne Edwards, 1864: Panopeidae
Ortmann, 1893; Palaeograpsus Bittner, 1875.
In this fossil association both species with burial habit
such as Lophoranina laevifrons (Bittner, 1875) and spe¬
cies typical of reefal environment such as Kromtìtis sp.
are present; this makes one presume that probably some
specimens were carried to thè deposition basin from thè
surrounding areas.
The discovery of Noetlingia veronensis (Bittner, 1 886)
that, after thè publication of thè initial work on thè spe¬
cies, had never been reported again and of “ Phrynolam¬
brus ” corallinus Bittner, 1 893 reported for thè first time
in thè Italian territory is remarkable.
The largest number of specimens discovered in this
deposit are representative of two species: Calappilia
dacica Bittner, 1893 and Kromtìtis sp.
Calappilia dacica, formerly found in upper Eocene
levels in Hungary (Bittner, 1893, Lòrenthey & Beurlen,
1929), had already been reported in “Main” quarry at
Arzignano (Vicenza) (Busulini et al., 1982); thè speci¬
mens found at Parona show thè spines on thè posterola-
teral margins to be smaller than those described by Bittner;
this peculiarity could be caused by different preservation.
In thè Eocene of Veneto, other species within thè same
genus have been found: C. gemmata Beschin, Busulini,
De Angeli & Tessier, 1994; C. incisa Bittner, 1886; and C.
subovata Beschin, Busulini, De Angeli & Tessier, 2002.
The genus Kromtìtis Miiller, 1984, also has previously
been recognized in Veneto: K. tetratuberculatus Beschin,
Busulini, De Angeli & Tessier, 2002, was found in “Main”
quarry at Arzignano while thè analysis of thè fossil fauna
discovered at Gecchelina, even though not complete,
highlights more than one species within thè same genus
(Beschin et al., 2000); thè Parona form differs from all
these and probably is a new species; such variety suggests
a suitable environment and rapid evolution of this group.
The discovery of Lobonotus cf. L. sandersi is particu¬
larly relevant: this species has been previously described
from thè Eocene of south Carolina within thè genus
Eohalimede (Blow & Manning, 1997) and subsequently
attributed to Lobonotus (Schweitzer et al., 2002): its pres¬
ence in northem Italy confirms thè well-known analogies
between these American faunas and those coeval ones in
Veneto (Feldmann et al., 1998, Beschin et al., 2002); if
thè generic attribution should be confirmed thè hypothesis
that thè geographic distribution of thè genus Lobonotus is
limited to thè Americas will have to be reconsidered.
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
13
Fig. 2 - A) Calappìlia dacica Bittner, 1 893 - MCZ 1 342. B) Noetlingia veronensis (Bittner, 1 886) - MCZ 1 343. C) Kromtitis sp. - MCZ
1346. D) “ Phrynolambrus ” corallinus Bittner, 1893 - MCZ 1345. E ) Lobonotus cf. L. sandersi - MCZ 1344. F) Palaeograpsus inflatus
Bittner, 1875 -MCZ 2522.
14
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
References
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1994 - I Crostacei eocenici della cava “Boschetto” di
Nogarole Vicentino (Vicenza - Italia Settentrionale).
Lavori - Società Veneziana di Scienze Naturali , Ve¬
nezia, 19: 159-215.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
2002 - Aggiornamento ai crostacei eocenici di cava
“Main” di Arzignano (Vicenza - Italia settentrionale)
(Crustacea, Decapoda). Studi e ricerche - Associazi¬
one Amici del Museo - Museo civico “G. Zannato ”,
Montecchio Maggiore (Vicenza): 7-28.
Beschin C., Busulini A., De Angeli A., Tessier G. &
Ungaro S., 2000 - The fauna of thè Gecchelina quarry
at Monte di Malo (Vicenza - Northern Italy): a prelim-
inary study. “lst Workshop on Mesozoic and Tertiary
decapod crustaceans ”, Studi e ricerche - Associazione
Amici del Museo - Museo civico ‘‘G. Zannato ”, Mon¬
tecchio Maggiore (Vicenza): 7-10.
Bittner A., 1886 - Neue Brachuyren des Eocans von
Verona. Sitzungsberichte der kaiserlichen Akademie
der Wissenschaften in Wien, Wien, 94 (I): 44-55.
Bittner A., 1893 - Decapoden des pannonischen Tertiàrs.
Sitzungsberichte der kaiserlichen Akademie der Wis¬
senschaften in Wien, Wien, 102 (II): 10-37.
Blow W. C. & Manning R. B., 1996 - Preliminary
descriptions of 25 new decapod crustaceans from thè
middle Eocene of thè Carolinas, U.S.A. Tulane Stud-
ies in Geology and Paleontology, New Orleans, 26
(1): 1-26.
Blow W. C. & Manning R. B., 1997 - A new genus,
Martinetto, and two new species of xanthoid crabs
from thè Middle Eocene Santee Limestone of South
Carolina. Tulane Studies in Geology and Paleontol¬
ogy, New Orleans, 30: 171-180.
Busulini A., Tessier G. & Visentin M., 1982 - Brachyura
della Cava Main (Arzignano) - Lessini orientali (Vi¬
cenza) (Crustacea, Decapoda). Lavori - Società Vene¬
ziana di Scienze Naturali, Venezia, 7: 75-84.
De Zanche V., Sorbini L. & Spagna V., 1977 - Geologia
del territorio del Comune di Verona. Memorie del
Museo Civico di Storia Naturale, Il serie, Verona, 1 :
1-51.
Feldmann R. M., Bice K. L., Hopkins C. S., Salva E. W.
& Pickford K., 1998 - Decapod crustaceans from thè
Eocene Castle Hayne Limestone, North Carolina:
paleoceanographic implications. The Paleontological
Society, Memoir 48, Supplement to Journal of Paleon¬
tology, Lawrence, 72 (1): 1-28.
Lòrenthey I. & Beurlen K., 1929 - Die fossilen Decapo¬
den der Lànder der Ungarischen Krone. Geologica
Hungarica, Serie Paleontologica, Budapest, 3: 1-420.
Schweitzer C. E., Feldmann R. M., Gonzàles-Barba G.
& Vega F. J., 2002 - New crabs from thè Eocene
and Oligocene of Baja California Sur, Mexico and
an assessment of thè evolutionary and paleobiogeo-
graphic implications of Mexican fossil decapods. The
Paleontological Society, Memoir 59, Supplement to
Journal of Paleontology, Lawrence, 76 (6): 1-43.
Claudio Beschin - Museo Civico “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: beschin.cl@libero.it
Alessandra Busulini - Via San Donà 160B, 30173 Venezia Mestre, Italy.
e-mail: busulini@tin.it
Giuliano Tessier - Via Barbarigo 10, 30126 Lido di Venezia, Italy.
e-mail: giultess@virgilio.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Gale A. Bishop
Mechanism for phosphatization of Dakoticancer assemblages
in thè Late Cretaceous western interior Seaway, USA
Fossil decapod crustaceans have often been described
as minor elements of molluscan assemblages and occa-
sionally been described from decapod-dominated assem¬
blages preserved as lagerstàtte. Because of their normal
scarcity in thè fossil record it is essential that maximal
information be consistently presented by authors that will
allow future workers to place thè decapod taxa into their
historical, evolutionary, ecological, and systematic con-
text. This has often been done by describing thè overall
aspects of a fauna; including its geographic, geologie, and
stratigraphic context.
Because of their complex exoskeleton, fossil decapods
are often incomplete and preserved as disassociation units
(Bishop, 1986; Bishop & Williams, 1986, 2005) that
normally disassociate as decomposition proceeds. Disas¬
sociation units typically consist of structural units of thè
exoskeleton, carapace, stemum, abdomen, and chelipeds
or pereiopods. Completeness of specimens varies from
assemblage to assemblage and depends upon thè physi-
cal, biological, and Chemical conditions of thè burial site
and diagenetic changes.
Lagerstatten are fossil assemblages that are remarkable
in either their diversity or quality of preservation, usuai ly
both. The Dakoticancer Assemblages in thè Pierre Shale
of south Dakota are largely represented by lagerstatten
(Fig. 1) whose structure and formative process have
been described, but whose formative genesis remained
unknown. The scenario envisioned (Bishop, 1981, 1986,
1987; Bishop & Williams 1986, 2005; Bishop et al.,
1998) involved periodic volcanic events that killed thè
phytoplankton and zooplankton in thè Cretaceous west¬
ern interior Seaway. The volcanic ejecta that rained onto
thè bottom enriching its nutrient load, which was then
exploited by a population explosion of segmented worms
which pelletized thè sediment, concentrating phosphate
in thè fecal pellets. This setting was than exploited by a
population explosion in decapods, which died and were
burrowed and pelleted by thè worms. The decapods
remains were somehow phosphatized by Chemical reac-
tions in thè sediment around thè buried decapods lying in
thè bentonitic substrate.
It is hypothesized herein that thè preservation of
these lagerstàtte is due to thè presence in thè substrate
of a sulfur and phosphate mediating bacterium related to
Thioploca, Beggiatoa, or Thiomargarita. Filamentous,
nitrate-accumulating sulfur bacteria described in 1907
of thè genus Thioploca Lauterbom, 1907, form exten-
sive populations of up to 120 g wet weight/m2 along thè
coast of Chile and Perù (1-3). Thiomargarita namibien-
sis described (Schultz et al., 1999), from thè Namibian
coast, is a colossal bacterium (nearly a millimeter in
diameter). Schulz & Schulz (2005: 416) demonstrated
that it accumulates intracellular polyphosphates under
aerobic conditions and releases phosphate under anoxic
conditions. These effects were measured into thè sub¬
strate with a maximum concentration at approximately
3 cm beneath thè water/sediment interface. This proc¬
ess creates pore water supersaturated in phosphate that
precipitates as phosphorite. The release of phosphate
by these organisms explains thè accumulation of many
phosphorites in thè geological record, including those
preserving thè Dakoticancer Assemblages. “Although
Thiomargarita appear to thrive best under low oxygen
or anoxic conditions, exposure to atmospheric oxygen
levels were not toxic as has been observed for Beggiatoa
(Nelson et al., 1986) and Thioploca (Maier & Gallardo,
1984)”.
Decapod assemblages described from north America
(Bishop, 1981, 1986, 1987; Bishop & Williams 1986;
Bishop et al., 1998) are decapod-dominated assemblages
overprinted on molluscan background assemblages. The
decapod assemblages from thè western interior are char-
acterized by: 1) being dominated by decapod crustaceans
numbering in thè thousands; 2) consisting of 1-3 dominant
taxa with 6-7 sub-dominant decapod taxa and numerous
non-decapod taxa; 3) being widespread over areas from a
few hundred to up to 1500 sq. km with discrete bounda-
ries; 4) being confìned to 1-3 m of bentonitic shale, but
not in discrete concentration layers; 5) being preserved
in phosphatic concretions exhibiting abundant burrows
and fecal pellets ascribable to segmented worms; 6) being
beautifully preserved, articulated decapods, often with
cuticle intact.
These assemblages have been interpreted (Bishop,
1981, 1986, 1987; Bishop & Williams 1986; Bishop et
al., 1998) as positive feedback systems involving vol¬
canic ash falls, which killed (directly or indirectly) thè
phyto- and zooplankton causing an enriching nutrient fall
onto thè shallow sea bottom. This initiated an explosion
of population growth in annelid worms and triggered a
population explosion of ubiquitous decapods in thè area
affected by and amenable to such population increases.
The burgeoning populations would continue to support
themselves over an interval of time, but would slowly
decline and tail off.
A thin-section of a phosphatic concretion containing a
transverse cross section of Dakoticancer overanus (Fig. 2)
shows carapace and pereiopods surrounded and filled by
burrowed and pelleted (muddy) sediment. Between and
around thè pellets and burrows is a diffuse field of small
crystallites. Comparison with data presented by Schultz
& Schultz, (2005; Fig. 3), shows similar crystallites of
phosphate within thè cells of Thiomargarita, surrounded
by vacuoles containing sulfur. The high concentration of
16
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
phosphate in sediment 3 cm beneath thè surface (Schultz
& Schultz, 2005), thè presence of phosphatic crystallites,
and thè burrowing by worms indicating shallow burial in
thè substrate are all consistent with thè proposed feedback
Fig. 1 - The crabs of thè Dakoticancer Lagerstàtten exhibit phenomenal
preservatimi, preserving significant portions of specimens, including
eye stalks, cuticle, and setal pits. Specimen enlarged approximately
two times; Late Cretaceous, (Maastrichtian), Pierre Shale, Mobridge
Member, Sitting Bull Locality, Corson County, south Dakota.
scenario of Bishop (1981, 1 986, 1 987) and Bishop & Wil¬
liams (1986).
It is suggested that sulphur and phosphate mediating
bacteria similar to Thiomargarita, if not ancestral to it,
were intimately involved in generating thè phosphate nec-
essary to form thè concretions preserving thè phosphatic
Dakoticancer lagerstàtten of south Dakota.
Fig. 2 - Transverse thin section of a concretion containing a complete
fossil crab, Dakoticancer overanus Rathbun, 1917. Carapace and legs
of thè crab are seen in cross section surrounded by large ovai fecal
pellets within and around thè crab, a worm burrow is present within
thè carapace and near thè stemum (not preserved), and open space
between pellets is filled with a clear minerai, possibly barite. The tiny
granular crystallites surrounding thè fecal pellets above thè carapace
and around thè circular worm burrow within thè carapace are thought
to be hydroxy-apatite grains derived from precipitation of polyphospate
inclusions similar to those figured by Schultz & Schultz, 2005 (Fig. 3).
References
Bishop G. A., 1981 - Occurrence and fossilization of thè
Dakoticancer Assemblage, Upper Cretaceous Pierre
Shale, South Dakota. In: Communities of thè Past, Gray
et al. (eds.), Hutchinson Ross Publishing Company.
Bishop G. A., 1986 - Taphonomy of thè North American
decapods. Journal of Crustacean Biology, 6 (3): 326-
355.
Bishop G. A., 1987 - Positive taphonomic feedback in
North American Tethyan Cretaceous decapod-worm
associations. In: Shallow Tethys 2. K. McKenzie (ed.).
Balkema Press, Rotterdam.
Bishop G. A., Feldmann R. M. & Vega F., 1998 - The
Dakoticancridae (Decapoda, Brachyura) from thè Late
Cretaceous of North America and Mexico. Contribu-
tions to Zoology, 67 (4): 237-255.
Bishop G. A. & Williams A. B., 1986 - The fossil
lobster Linuparus canadensis. Cadile Shale (Creta¬
ceous, Turonian), Black Hills. National Geographic
Research, 2 (3): 372-387.
Bishop G. A. & Williams A. B., 2005 - Taphonomy and
Preservation of Burrowing Thalassinidean Shrimps.
Proceedings of thè Biological Society of Washington,
Washington, 118 (1): 218-236.
Maier S. & Gallardo V. A., 1984 - Thioploca araucae
sp. nov. and Thioploca chileae sp. nov. International
Journal of Systematic Bacteriolology, 34: 414.
Nelson D. C., Revsbech N. P. & Jorgensen B. B., 1986 -
Microoxic-anoxic niche of Beggiatoa spp.: microelec-
trode survey of marine and ffesh-water strains. Applied
and Environmental Microbiology, 52: 161-168.
Schulz H. N., Brinkhoff T., Ferdelman T. G., Flemandez
M., Teske M. A. & Jorgensen B. B., 1999 - Dense
Populations of a Giant Sulfur Bacterium in Namibian
Shelf Sediments. Science, 284: 493-495.
Schultz H. N. & Schultz H. D., 2005 - Large Sulfur Bac¬
teria and thè Formation of Phosphorite. Science, 307 :
416-418.
Gale A. Bishop - Emeritus Professor of Geology, Department of Geology and Geography, Georgia Southern University,
Statesboro, Georgia 30460, U.S.A.
e-mail: gbishop@geotrec.org
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Gérard Breton & Joe S. H. Collins
Decapod fauna from thè Cenomanian stratotype
During thè 1 9th century, thè stratotypic Cenomanian,
i.e. thè Sarthe department in France, around thè town
of Le Mans, provided locai collectors such as J. Triger
and E. Guéranger (Breton, 1996) a large amount of well
preserved fossils. Among them, decapods in particular
were collected from thè so-called “Couche à Crastacés”
in thè “carrière de la Butte de Gazonfier” within thè town
of Le Mans. The “Couche à Crustacés” is located in thè
first lowest meters of thè Sables du Perche Formation,
Middle-Upper Cenomanian, Jukesbrownei-Guerangeri
biozones. This quarry (section in Guinot & Breton,
2006) has been disused since thè early 20th century. Other
formations in thè stratotype area provided decapods:
Lower Cenomanian, Mames de Ballon Formation,
Carcitanensis and base of Saxbii biozones; Sables &
Grès de Lamnay Formation, around Lamnay and St.
Maixent, limit of thè Mantelli and Dixoni biozones.
Owing to their equal richness in crustaceans, Triger (in
de Hennezel, 1858) erroneously correlated thè Mames
de Ballon with thè Couche à Cmstacés in thè Sables
du Perche; in thè museum collections, thè two origins
are often mixed. For thè geological context and history,
see Guinot & Breton (2006); for thè stratigraphical
correlations, see Juignet (1974, 1980).
Because thè Butte quarry is now disused, fresh
decapod material is rare, and most of thè specimens noted
here come from museum collections: Muséum d’Histoire
naturelle, Le Havre (MHNH); Musée d’Histoire natu-
relle Le Mans (register numbers MHN LM 2003-1-
XXXX abbreviated here LMXXXX); Muséum national
d’Histoire naturelle, Paris, Domaine des Sciences de la
Terre (MNHN); Laboratoire de Géologie, Université de
Caen (LGUC); Université Lyon I (FSL); Université de
Paris VI (UPMC).
Principal publications concerning thè study of these
Cenomanian decapods include works of A. Milne
Edwards (1861, 1862a, 1862b, 1882), A. Milne Edwards
& Brocchi (1879), Brocchi (1887), Van Straelen (1936),
and Guinot & Breton (2006).
The most frequently encountered macruran is
Hoploparia trigeri (Van Straelen 1936) of which we
here designate as thè lectotype thè specimen figured
as “topotype” by Van Straelen (1936, pi. 3, fig. 4)
[LM3760] from La Butte quarry. Other specimens are:
LM3799A and MNHN A25925 (Fig. 1) with elongated
chelipedes (first known occurrence of associated
chelae), LM3801 juvenile, LM3785 has an irregular
bopyriform swelling, UPMC unregistered [from La
Butte quarry], MNHN A25926, [from St-Mars-sous-
Ballon].
The palinurid Linuparus dentatus Van Straelen,
1936 [ non 1931] is far less frequent. The 1931 paper
of Van Straelen has not been found, either in thè Van
Straelen’s library in thè MNHN, Paris, or in Bruxelles
(J.-M. Bragard, pers. comm.) and it is not quoted in his
bibliographic list (Corin, 1965): thè citation of this 193 1
paper is supposed to be erroneous, thus we retain thè
date, 1936, for this species. We designate as lectotype
thè specimen figured by Van Straelen (1936, pi. I, fig.
2) [MNHN R03384]. A plaster cast of thè lectotype
[LM3781] and fragmentary specimens [LM3789].
Callianassa cenomaniensis A. Milne Edwards,
1861. We have not been able to fìnd thè specimen(s)
drawn by A. Milne Edwards (1861, pi. 14, fig. 5, 5A),
therefore we designate as lectotype thè specimen fig¬
ured here Fig. 2 [MNHN R03548], which fits well thè
originai description. It is labelled “1902-3” like other
specimens studied by A. Milne Edwards and it Comes
from thè type locality: St-Mars-sous-Ballon, Lower
Cenomanian. The matrix, a fine glauconitic limestone
with one Orbitolina concava (Lamarck 1816) proves it
comes from thè Marnes de Ballon Formation. This spe¬
cies is fairly abundant throughout thè stratotypic Ceno¬
manian and there are numerous specimens in MHN LM,
MNHN, FSL, mainly from Marnes de Ballon, but also
from Sables du Perche Formation.
Porcellana antiqua A. Milne Edwards, 1882 is con-
sidered as a nomen dubium because thè originai descrip¬
tion is based on a single specimen, not illustrated, which
has not been found.
Pagurus sp.? [MNHN B16584B]. A small hermit
crab is preserved in thè aperture of an unidentifìed,
diagenetically compressed gastropod shell. It displays
fragmentary appendages: thè incomplete punctuated right
chela seems to be larger than thè left one (Fig. 3). Such
preservation is rare (Jagt et al., 2006; Todd & Collins,
2005). Labelled “Crustacé de la Sarthe communiqué
par M. Triger 1902-3. Crétacé sup”, it probably comes
from thè Carrière de la Butte, but not from thè Couche à
Crustacés. The matrix, a grey soft clay with ferrugineous
staining, and thè type of fossilization with a complete
decalcification evoke thè “Lentilles à Echinodermes”,
10 meters or so below thè Couche à Crustacés, within
thè Sables & Grès du Mans Formation (Breton, 1996;
Guinot & Breton, 2006: fig. 1).
Caloxanthus formosus A. Milne Edwards, 1862.
There are a few specimens, of both carapaces and
chelipeds. No ventral side is known. [LM3792
(carapace); LM3796, 3797 (chelipeds); MNHN
R03351 (carapace), B 16572 (carapace and associated
chelipeds)].
Trachynotocarcinus sulcatus (Bell, 1863). One
incomplete small carapace [MNHN B 16571], Upper
Cenomanian, Saint-Calais (Sarthe).
18
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Raninella trigeri A. Milne Edwards, 1862. Type
species of thè genus Raninella, by originai designation.
Although this species, and thè following one are both
named by A. Milne Edwards (1862b), and adequately
distinguished in barely two lines, this sufficiently
constitutes a definition of both taxa in respect to art.
12.1 of ICZN. “this, though jejune, is just adequate to
make thè name available” (Wright & Collins, 1972).
We designate as thè lectotype thè specimen MNHN
B 16565, figured by Brocchi (1877, pi. 29, figs. 1,
2). Paralectotype MNHN R0394 figured by Brocchi
(1877, pi. 29, fig. 3). Other specimens and casts of
type material in MHN LM. Their matrix shows that all
specimens come from thè Carrière de la Butte, Couche
à Crustacés.
Raninella elongata A. Milne Edwards, 1862. The
differences with R. trigeri are well explained by Broc¬
chi (1877). Raninella elongata is always smaller than
R. trigeri, and we have not seen any intermediate sized
Raninella in thè collections we visited. Thus we do not
agree with Van Straelen (1936) who stated that R. elon¬
gata and R. trigeri are synonymous, thè former being a
juvenile of thè latter. Type series in MNHN, formerly
B 16575 with 5 specimens from Sables du Perche Fm,
La Butte quarry and 4 specimens from Marnes de
Ballon Fm, with a single label “Crustacés des Grès
verts du Mans A. Milne Edwards 1902-3”. Among
them, we designate as lectotype MNHN A25922 thè
specimen figured by Brocchi (1877, pi. 29, figs. 4, 5),
La Butte quarry. Paralectotypes are: MNHN A25923,
figured pi. 29, fig. 5 bis, Ballon; 3 specimens MNHN
B 16575, La Butte; 4 specimens MNHN A25924,
Ballon; other paralectotypes: 2 specimens labelled
“Raninella Trigeri A. M. Edw. Cénomanien Le Mans
Sarthe 1902-3”, B 16564, La Butte quarry. Other speci¬
mens LM3788.
Raninidae gen. nov., sp. nov. from thè Lower
Cenomanian Sables & Grès de Lamnay Fm, St-Maix-
ent (Sarthe), two specimens that will be described and
named later: Collins & Breton (in progress).
Cenomanocarcinus injìatus Van Straelen, 1936.
Figured as Necrocarcinus inflatus ( nomen nudum:
ICZN 13.1.1) by Boule & Piveteau (1935), fig. 670.
We designate as lectotype thè specimen figured by Van
Straelen (1936, pi. IV, fig. 8) [MNHN J08587 ex coll.
Hébert, labelled “La Butte de Gazonfier au Mans”].
Other material: MNHN R05504 (paralectotype),
MNHN B16588a: anterior fragment of thè carapace,
with associated left cheliped (Fig. 4), LM3780, 3805,
3806, a juvenile with produced epibranchial, postero-
laterally directed, spines which, in combination with
thè triple row of tubercles, gives it a calappid-like
appearance (Fig. 5). These spines are broken in all
other specimens, giving them a more rounded outline.
Necrocarcinus labeschii (Deslongchamps, 1835).
One specimen (carapace length 19mm; width: 24mm),
Sables du Perche Formation, Le Mans [LM3808, coll.
Guéranger]. Two small cephalothoraces [one at least
presumably from Sables du Perche Fm], ca 12 x 12mm
[LM3807] are labelled “ Necrocarcinus minutus M.
Edw.”. Necrocarcinus minutus is a nomen nudum in
Guillier (1886). These two specimens seem to be juve-
niles of N. labeschii (Fig. 6).
Lithophylax trigeri A. Milne Edwards & Brocchi,
1879. Lectotype MNHN A25835 selected among thè
type series by Guinot & Breton (2006). For a com¬
plete description, see Guinot & Breton (2006). This
small crab, which has a stridulating apparatus and was
coprophagic (Breton, 2006), is by far thè most abun-
dant decapod from thè stratotypic Cenomanian: many
specimens in MNHN, MHN LM, MHNH, LGUC,
Muséum de Rouen: coll. Fortin 31217, Naturai His-
tory Museum in Bruxelles (B), and thè Naturai History
Museum (UK).
Xanthosia sp.? One specimen, without any indica-
tion of origin, but within thè Guéranger collection
[LM3798] may come from thè Sables du Perche For¬
mation.
The decapod fauna from thè Cenomanian stratotype
is varied, and contains apparently endemie species.
Among thè 14 taxa quoted here, 10 were found in
thè Carrière de la Butte, Sables du Perche Formation,
Couche à Crustacés. But Lithophylax trigeri is also
found in thè Cenomanian from thè Aude department,
south France (Van Bakel, Artal, Breton, Guinot &
Vizcaino, in progress). Trachynotocarcinus sulcatus
is also known from England (Wright & Collins, 1972)
and from Normandy (Breton, unpublished). Caloxan-
thus formosus is also found in Normandy (Breton,
unpublished) and in thè south of France (Van Straelen,
1938). Callianassa cenomaniensis was collected in
Normandy, and Raninella trigeri and Cenomanocarci¬
nus inflatus were found in thè south of France (Van
Straelen, 1936, 1938). Necrocarcinus labeschii is
widespread in England and France (Wright & Collins,
1972). The other species seem to be restricted to Le
Mans area, but they are indeed thè rarest.
The apparent endemism among stratotypic Ceno¬
manian decapod fauna is undoubtedly due to thè pecu-
liar palaeoenvironmental and taphonomic conditions
which led to thè “Couches à Crustacés” Konservat-
Lagerstàtte. As in thè case of recently discovered
Lithophylax trigeri outside thè type area mentioned
above, subsequent collecting could well produce a
wider distribution of “pseudo-endemie” species.
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
19
Fig. 1 - Hoploparia trigeri (Van Straelen, 1936). MNHN A25925.
Middle - Upper Cenomanian, Sables du Perche Formation, “Couches
à Crustacés” beds, La Butte quarry, Le Mans (Sarthe, France). Moult
with right cheliped. Scale bar: 1 cm.
Fig. 2 - Callianassa cenomaniensis A. Milne Edwards, 1861. Lecto-
type, MNHN R03548. Lower Cenomanian, Mames de Ballon Forma¬
tion, Saint-Mars-sous-Ballon (Sarthe, France). Scale bar: 1 cm.
Fig. 3 - Pagurus sp.? MNHN B16584B. Probably Middle - Upper
Cenomanian, Sables & Grès du Mans Formation, “Couches à Echino-
dermes” beds, La Butte quarry, Le Mans (Sarthe, France). Scale bar:
5 mm.
Fig. 4 - Cenomanocarcinus inflatus Van Straelen, 1936. Paralectoptype,
MHNH B16588a. Probably Middle - Upper Cenomanian, Sables du
Perche Formation, “Couches à Crustacés” beds, La Butte quarry, Le
Mans (Sarthe, France). Cheliped. Scale bar: 1 cm.
Fig. 5 - Cenomanocarcinus inflatus Van Straelen, 1936. LM 3806, coll.
Guéranger. Middle - Upper Cenomanian, Sables du Perche Forma¬
tion, “Couches à Crustacés” beds, La Butte quarry. Le Mans (Sarthe,
France). Carapace of a young individuai with pronounced epibranchial
spines. Scale bar: 1 cm.
Fig. 6 - Necrocarcinus labeschii (Deslongchamps, 1835). LM 3807,
coll. Guéranger, labelled as “ Necrocarcinus minutus M. Edw. "[nomen
nudum in Guillier (1886)]. Probably Lower Cenomanian, Mames de
Ballon Formation, Ballon (Sarthe, France). Carapace of a very young
individuai. Scale bar: 5 mm.
20
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
References
Boule M. & Piveteau J., 1935 - Les Fossiles - Eléments
de paléontologie. Masson & Cie, Paris.
Breton G., 1996 - Tethyaster guerangeri sp. nov. (Astro-
pectinidae, Asteroidea, Echinodermata): deux spéci-
mens d’astérides d’une conservation exceptionnelle
du Cénomanien du Mans (Sarthe, France). Bulletìn
de la Société Géologique de Normandie et des Amis
du Muséum du Havre, Le Havre, 82 (4), 1995 (1996):
17-29.
Breton G., 2006 - Un cas de paléocoprophagie chez un
crabe du Cénomanien du Mans (collection Boutillier,
Université de Caen). Bulletìn de la Société linnéeenne
de Normandie, Caen, 119: 41-43.
Brocchi P., 1 877 - Description de quelques Crustacés fos¬
siles appartenant à la tribù des Raniniens. Annales des
Sciences Géologique, Paris, 8: 1-8.
Corin F., 1965 - Victor Van Straelen (14 juin 1889 - 29
février 1964) [Notices nécrologiques]. Bulletin de
la Société Belge de Géologie, de Paléontologie et
d’Hydrologie, Bruxelles, 74: 12-35.
Guillier A., 1886 - Géologie du Département de la Sarthe.
Monnoyer, Le Mans.
Guinot D. & Breton G., 2006 - Lithophylax trigeri A.
Milne-Edwards & Brocchi, 1879 from thè French Cre-
taceous (Cenomanian) and placement of family Litho-
phylacidae Van Straelen, 1936 (Crustacea, Decapoda,
Brachyura). Geodiversitas, Paris, 28 (4): 591- 633.
Jagt J. W. M., van Bakel B. W. M., Fraaije R. H. B.
& Neumann C., 2006 - In situ fossil hermit crabs
(Paguroidea) from northwest Europe and Russia.
Preliminary data on new records. Revista Mexicana de
Ciencias Geológicas, Mexico, 23, 3: 464-469.
Juignet P., 1974 - La transgression crétacée sur la bor¬
dure orientale du Massif armoricain. Aptien, Albien,
Cénomanien de Normandie et du Maine. Le stratotype
du Cénomanien. Thèse de Doctorat d’Etat, Université
de Caen (unpublished).
Juignet P., 1980 - Cénomanien. In: Synthèse géologique
du Bassin de Paris. Volume I. Statigraphie et
paléogéographie. Mégnien C. & Mégnien F. (eds.).
Mémoire BRGM, Paris, 101: 292-297.
Milne Edwards A., 1861 - Histoire des Crustacés Podo-
phtalmaires fossiles. Victor Masson & fils , Paris, 1.
Milne Edwards A., 1862a - Monographie des Crustacés
fossiles de la famille des Cancériens. Première Partie.
Annales des Sciences Naturelles (Zoologie), Paris, 4
(18): 31-85.
Milne Edwards A., 1862b - Sur l’existence de Crus¬
tacés de la famille des Raniniens pendant la péri-
ode crétacée. Comptes rendus hebdomadaires des
Séances de l’Académie des Science , Paris, 55: 492-
494.
Milne Edwards A., 1 882 - Note sur un Crustacé du terrain
crétacé appartenant au genre Porcellana. Annales de
Sciences géologiques, Paris, 12: 52 [with a mistake in
thè pagination: p. 42 inserted at a wrong place in thè
same fascicle]
Milne Edwards A. & Brocchi P., 1 879 - Note sur quelques
Crustacés fossiles appartenant au groupe des Macro-
phthalmiens. Bulletin de la Société Philomatique de
Paris, Paris, 7 (3): 113-117.
Van Straelen V., 1936 - Crustacés Décapodes nouveaux
ou peu connus de Fépoque crétacique. Bulletin du
Musée royal d’ Histoire naturelle de Belgique, Brux¬
elles, 12 (45): 1-50.
Van Straelen V., 1938 - La faune carcinologique du Cré¬
tacique des Corbières méridionales. Compte Rendu
sommaire des Séances de la Société géologique de
France, Paris, 1938 (9): 151-153.
Todd J. & Collins J. S. H., 2005 - Neogene and Qua-
temary crabs (Crustacea, Decapoda) collected from
Costa Rica and Panama by members of thè Panama
Palaeontological project. Bulletin of thè Mizunami
Fossil Museum, Mizunami, 32: 53-85.
Triger J., 1858 - Note sur la composition du terrain
crétacé du département de la Sarthe In: de Hennezel
(ed.). Bulletin de la Société d’Agriculture, Sciences et
Arts de la Sarthe, Le Mans, 2 (13): 201-211.
Wright C. W. & Collins J. S. H., 1972 - British Cretaceous
crabs. Palaeontographical Society ( Monographs ),
London, 533: 1-114.
Gérard Breton - Chercheur libre rattaché au Laboratoire Géosciences, Université Rennes I, 35042 Rennes Cedex, France.
rue des Réservoirs, 76600 Le Havre, France.
e-mail: gerardbreton@free.fr
Joe S. H. Collins - 8, Shaw’s Cottages. Perry Vale, London SE23 2QN, UK.
Department of Palaeontology, Naturai History Museum, London SW7 5BD, UK.
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Antonio De Angeli & Claudio Beschin
Tertiary stomatopods from Italy
De Angeli & Garassino (2006) reported Decapoda
and Stomatopoda from thè Triassic to Pleistocene of
Italy, pointing out that thè stomatopods are very
rare and usually incomplete in thè fossil record. De
Angeli & Beschin (2006) described two new species
of stomatopods, giving a check list of Tertiary spe¬
cies from Veneto. The Tertiary species known to date
are thè following: Pseudosquilla berica De Angeli &
Messina, 1996 (lower Oligocene, Perarolo - Vicen¬
za) (Pseudosquillidae Manning, 1977); Lysiosquilla
antiqua (Munster, 1842) (lower-middle Eocene,
Monte Bolca - Verona), Lysiosquilla messinai De
Angeli, 1997 (lower Oligocene, Perarolo - Vicenza),
Lysiosquilla sp. (middle Eocene, Laverda - Vicenza)
(Lysiosquillidae Giesbrecht, 1910); Coronidopsis
albanellensis De Angeli & Beschin, 2006 (middle
Eocene, Chiampo Valley - Vicenza) (Eurysquillidae
Manning, 1977); Squilla miocenica Lovisato, 1894
(Miocene, Cagliari - Sardinia), Squilla breoniensis
De Angeli & Beschin, 2006 (middle Eocene, Breonio
- Verona) (Squillidae Latreille, 1802).
Pseudosquilla berica De Angeli & Messina,
1996 (lower Oligocene, Perarolo - Vicenza) has a
convex carapace transversely marked by two lon-
gitudinal grooves, rostrum with rounded margin,
thoracic somites smooth, abdominal somites III- V
marked by a longitudinal groove, abdominal somite
V with posterior carina, abdominal somite VI with
four small posterior spines, and telson with median
carina and six marginai teeth. The shapes of thè right
exopod and endopod, raptorial claw, and eyestalk are
known, too.
Lysiosquilla antiqua (Mtinster, 1842) (lower-
middle Eocene, Monte Bolca) was described on thè
morphological characters of thè holotype which is
lost. Later, Secretan (1975) gave a new description
of this species based upon many specimens, housed
in thè Museo di Storia Naturale di Verona and
Museo di Geologia e Paleontologia dell’Università
di Padova. Schram & Miiller (2004) considered L.
antiqua minor Secretan, 1975, to be a synonym with
L. antiqua (Miinster, 1842).
Lysiosquilla messinai De Angeli, 1997 (lower
Oligocene, Perarolo - Vicenza) has a carapace
longer than wide, pentagonal rostrum, abdominal
somites with curved margin, subrectangular and fiat
telson with weak median carina and small marginai
teeth. Besides thè stomatopods, anomurans ( Upoge -
bia perarolensis De Angeli & Messina, 1992, and
Ga/athea valmaranensis De Angeli & Garassino,
2002), isopods ( Cirolana fabianii De Angeli &
Rossi, 2006) and mysidaceans ( Mysidopsis o/igoce-
nicus De Angeli & Rossi, 2006) were described from
Perarolo (De Angeli & Messina, 1992, 1996, 1997;
De Angeli, 1997; De Angeli & Rossi, 2006).
Lysiosquilla sp. (lower Lutetian, Laverda - Vi¬
cenza) was described from a single dactylus of thè
raptorial claw. De Angeli & Beschin (2006) also
reported thè brachyurans Harpactocarcinus punc-
tulatus (Desmarest) and Palaeograpsus inflatus Bit-
tner from this locality.
Coronidopsis albanellensis De Angeli & Beschin,
2006 (middle Eocene of “Albanello” quarry, Noga-
role Vicentino - Vicenza) has abdominal somites
subrectangular and smooth, with a latero-posterior
pleural widening, telson with two weak subme-
dian carinae, raptorial claws with elongate pro-
podus having three spines, and dactylus with three
strong spines, besides thè elongate distai spine.
Beschin et al. (2005) reported one dactylus from
Grola di Comedo Vicentino (Vicenza) beloning to
C. albanellensis. Coronidopsis albanellensis was
ascribed to Coronidopsis for thè abdomen, smooth
telson without carinae, and four spines of dactylus
of thè raptorial claw. Coronidopsis is known from
two Recent species C. bicuspis Hansen, 1926, and
C. serenei Moosa, 1973, widespread in Indo-Pacific
area, New Caledonia, and Australia.
Squilla breoniensis De Angeli & Beschin, 2006
(middle Eocene, Breonio - Verona) has an elongate
carapace with marked cervical groove, and short
posterior median ridge, abdominal somites with
a pair of submedian, intermedian, and lateral
ridges, telson with median carina and serrate lateral
margins, raptorial claws with propodus having
small spines along thè margins, and a dactylus with
four-five strong spines. Squilla is known from thè
Cretaceous to thè Recent with seven fossil species,
S. breoniensis, S. cretacea Schliiter, 1868, S.
hollandi Forster, 1982, S. laingae Hof & Schram,
1998, S. miocenica Lovisato, 1894, S. sonomana
(Rathbun, 1926), and Squilla sp., and 23 Recent
species (Schram & Miiller, 2004).
Squilla miocenica Lovisato, 1894 (Miocene, Fan-
gario - Sardinia) was described by Lovisato (1894,
figs. 1-3) on morphological characters of some dac-
tyli with eight spines (including thè elongate spine).
Via Boada (1949) also reported this species from thè
Miocene of Spain.
Six species known to date in Italy come from
Veneto, attesting to thè importance of thè Eocene
and Oligocene quarries of this area, very rich of
decapods, useful to establish a relationship between
thè fossil and Recent taxa.
22
ED1TED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Fig. 1 - A-B) Squilla breoniensis De Angeli & Beschin, 2006, holotype, n. cat. MCZ 1552 - I.G. 284630. A) Lateral view. B) Dorsal
view. C-E) Coronidopsis albanellensis De Angeli & Beschin, 2006, holotype, n. cat. MCZ 1542 - I.G. 284620. C) Dorsal view. D)
Lateral view. E) Raptorial claw.
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
23
References
Beschin C., De Angeli A., Checchi A. & Zarantonello
G., 2005 - Crostacei eocenici di Grola presso Spag-
nago (Vicenza, Italia settentrionale). Studi e Ricerche
- Associazione Amici del Museo - Museo Civico “G.
Zannato”, Montecchio Maggiore (Vicenza), 12: 5-35.
De Angeli A., 1997 - Lysiosquilla messinae, nuova specie
di crostaceo stomatopode del Terziario di Vicenza
(Nord Italia). Studi e Ricerche - Associazione Amici
del Museo - Museo Civico “G. Zannato ”, Montecchio
Maggiore (Vicenza), 1997: 23-26.
De Angeli A. & Beschin C., 2006 - Stomatopodi terziari
del Veneto (Italia settentrionale). Studi e Ricerche -
Associazione Amici del Museo - Museo Civico “G.
Zannato”, Montecchio Maggiore (Vicenza), 13: 19-
28.
De Angeli A. & Garassino A., 2006 - Catalog and bibli-
ography of thè fossil Stomatopoda and Decapoda from
Italy. Memorie della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale di Milano ,
Milano, 35 (1): 1-95.
De Angeli A. & Messina V., 1992 - Upogebia perarolen-
sis nuova specie di crostaceo del Terziario del Veneto
(Italia). Lavori - Società Veneziana di Scienze Natu¬
rali, Venezia, 17: 183-191.
De Angeli A. & Messina V., 1996 - Pseudosquilla
berica nuova specie di Stomatopoda del Terziario
Veneto. Studi e Ricerche - Associazione Amici del
Museo - Museo Civico “G. Zannato”, Montecchio
Maggiore (Vicenza), 1996: 5-10.
De Angeli A. & Messina V., 1997 - Galathea xvein-
furteri Bachmayer, 1950 (Crustacea, Anomura)
nell’Oligocene di Perarolo (Vicenza, Nord Italia).
Studi e Ricerche - Associazione Amici del Museo -
Museo Civico “G. Zannato”, Montecchio Maggiore
(Vicenza), 1997: 17-21.
De Angeli A. & Rossi A., 2006 - Crostacei oligocenici
di Perarolo (Vicenza - Italia settentrionale), con la
descrizione di una nuova specie di Mysida e di Iso-
poda. Lavori - Società Veneziana di Scienze Naturali,
Venezia, 31: 85-93.
Lovisato D., 1894 - Avanzi di Squilla nel Miocene medio
di Sardegna. Rendiconti della Reale Accademia dei
Lincei, Roma, III, 4: 205-209.
Schram F. R. & Miiller H-G., 2004 - Catalog and bibliog-
raphy of thè fossil and recent Stomatopoda. Backhuys
publishers, Leiden, The Netherlands.
Secretan S., 1975 - Les Crustacés du Monte Bolca. Museo
Civico di Storia Naturale di Verona - Studi e Ricerche
sui giacimenti terziari di Bolca, Verona, 2: 315-425.
Via Boada L., 1949 - Un resto de estomatópodo {Squilla
miocenica Lovisato) en las margas tortonienses de
Montjuich. Bulletin de la Istitució Catalana Història
Naturai, Barcellona, 37: 2-4.
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
Claudio Beschin - Museo Civico ‘‘G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: beschin.cl@libero.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Antonio De Angeli & Alessandro Garassino
Decapod crustaceans from thè Mesozoic and Cenozoic
of Friuli-Venezia Giulia (NE Italy)
The study of brachyurans and anomurans from thè
Mesozoic and Cenozoic of Friuli-Venezia Giulia (NE
Italy) was previously limited to only two reports by
Dainelli (1915), and Collins & Dieni (1995). Dainelli
(1915) reported four incomplete specimens of Lopho-
ranina maresticma (Kònig, 1825) from thè Eocene of
Buttrio and Valle del Natisone (Udine), two specimens
of Palaeocarpilius macrochelus (Desmarest, 1 822) from
Monte Plauris (Udine), and one specimen of Panopeus
vicentinus Bittner, 1875, from Buttrio. Collins and Dieni
(1995) reported one incomplete chela of indeterminate
pagurid and one incomplete carapace ascribed to Grapto-
carcinus bellona Collins & Dieni, 1995, from thè Upper
Cretaceous (Cenomanian) of Col dei Schiosi (Altopiano
del Cansiglio, Pordenone).
The recent discovery of thalassinids, anomurans, and
brachyurans from some localities, Altopiano Prat, Casali
Ottelio, and Borgo Vigant (Udine province), Val Caltea,
Monte Ciaurlec, Almadis, and Meduno (Pordenone prov¬
ince) permitted increasing of thè carcinologie knowledge
about these systematic groups (De Angeli & Garassino,
2006).
The studied sample included Corallianassa rigoi
De Angeli & Garasino, 2006 (infraorder Thalassinidea
Latreille, 1831), Calteagalathea friulana De Angeli &
Garassino, 2006 (infraorder Anomura MacLeay, 1838),
Pithonoton margìnatum (v. Meyer, 1842), Graptocarci-
nus bellonii Collins & Dieni, 1995, Lophoranina mares-
tiana (Kònig, 1825), Portunus monspeliensis (A. Milne
Edwards, 1860), Portunites rosenfeldi De Angeli & Ga¬
rassino, 2006, and Titanocarcinus raulinianus A. Milne
Edwards, 1 863 (infraorder Brachyura Latreille, 1 802).
Some chelae were ascribed to Corallianassa
rigoi from thè Cretaceous (Aptian-Campanian)
of Monte Ciaurlec (Pordenone). The oldest fossil
species belonging to this genus is C. acucur-
vata Swen, Fraaije & Van der Zwaan, 2001, from
thè Upper Cretaceous (Maastrichtian) of Maas¬
tricht (The Netherlands) (Swen et al., 2001). Co
rallianassa acucurvata differs from C. rigoi by thè
smooth lower margin of thè bigger chela, and by thè
fìxed and movable fingers being more elongate. Co¬
rallianassa sp. from thè Paleocene (upper Danian)
of Copenhagen (Denmark), described by Rasmussen
(1971) is also assigned to this genus. The Danish spe¬
cies differs from C. rigoi by thè bigger chela with more
sinuous lower margin, and by thè fìxed and movable
fingers being more developed. The discovery of C.
rigoi is very important because it represents thè first
report of this genus in Italy, attesting to thè widespread
distribution of Corallianassa in thè European seas
during thè Upper Cretaceous.
Calteagalathea friulana , from thè Upper Creta¬
ceous (Maastrichtian) of Val Caltea (Pordenone),
represents thè oldest galatheid known to date in Italy,
showing morphological affinities with thè representa-
tives of Paragalathea, which is widespread in thè
Upper Jurassic and Cretaceous of Europe. A compari-
son between C. friulana and some pictures of thè holo-
type of P. multisquamata Via Boada, 1981, housed in
thè Seminary Museum of Barcelona, suggested that thè
Spanish species had a wider and more curved cervical
groove, smaller, thicker, and more uniform granula-
tion, more rectilinear lateral margins, and less deep
postcervical depression.
The discovery of Pithonoton marginatum (v.
Meyer, 1842) from thè Upper Jurassic (Oxfordian-
Kimmeridgian) of Altopiano Prat (Udine) was very
interesting since this prosopid represents thè oldest
brachyuran known to date in Italy. The discovery of
prosopids in Italy was until now limited to thè Titho-
nian of Sicily (Gemmellaro, 1869) with Pithonoton
bidentatum (Reuss, 1859) (= Prosopon etalloni Gem¬
mellaro in Gemmellaro, 1869), and Pithonoton grande
(v. Meyer, 1860) (= Prosopon polyphemi Gemmellaro
in Gemmellaro, 1869). The report of prosopids from
thè Upper Jurassic of Friuli-Venezia Giulia extended
thè geographical distribution of these decapods also
into N Italy.
The discovery of one well-preserved specimen of
Graptocarcinus bellonii attested to thè spread of this
dynomenid in thè Upper Cretaceous (Maastrichtian)
of Friuli-Venezia Giulia, species already described
by Collins & Dieni (1995), from thè Upper Creta¬
ceous (Cenomanian) of Col dei Schiosi (Altopiano del
Cansiglio, Pordenone). However, a comparison with
thè holotype (DGP. B241), housed in thè Museo di
Geologia e Paleontologia dell’Università degli Studi
di Padova, resulted in thè following observations: thè
studied specimen is more complete than thè holotype,
lacking only thè fronto-orbital and anterolateral mar¬
gins; thè dorsal ornamentation of thè studied specimen
is less uniform; thè branchial, hepatic, and gastric
regions of thè studied specimen have smaller tubercles,
while thè tubercles are rare in thè median part. The
different ornamentation of thè surface of thè carapace
between thè holotype and thè studied specimen could
be related to different onthogenetic stages since thè
carapace of thè studied specimen is three time larger
than that of thè holotype.
The discovery of new specimens of Lophoranina
marestiana (Kònig, 1825) attested to thè widespread
distribution of this species in thè Eocene of Friuli-
Venezia Giulia, where it was already known by four
3 1(0 SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
25
Fig. 1 - A) Corallianassa rigoi De Angeli & Garassino, 2006. B) Calteagalathea friulana De Angeli & Garassino, 2006, holotype.
C) Pithonoton marginatimi (v. Meyer, 1 842). D) Portunites rosenfeldi De Angeli & Garassino, 2006. E) Graptocarcinus bellona Col¬
lins & Dieni, 1995. F) Titanocarcinus raulinianus A. Milne Edwards, 1863.
26
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
incomplete carapaces, discovered in Buttrio and
Valle del Natisone by Teliini and reported by Dainelli
(1915). Lophoranina marestiana is particularly wide-
spread in thè Cuisian and Lutetian levels of Veneto
(Beschin et al., 1988).
The discovery of some specimens of Portu-
nus monspeliensis (A. Milne Edwards, 1860) (= P.
granulatus A. Milne Edwards, 1860) from thè middle
Miocene (Langhian) of Meduno (Pordenone) is impor-
tant because this species is reported for thè first time
in Friuli-Venezia Giulia and its presence attests to
thè widespread distribution of this portunid in thè
Mediterranean Sea during thè Miocene. This species
is known from thè Miocene of S Europe (Hungary,
France, Austria) and N Africa (Egypt), while Ristori
(1888), Lòrenthey (1909), Comaschi Caria (1950,
1956), and Marras & Ventura (1991) reported this spe¬
cies in Italy from thè Miocene of Sardinia and Emilia
Romagna. The presence of P. monspeliensis from thè
Rupelian (lower Oligocene) of Bacino Ligure Pie¬
montese is dubious. In fact, Allasinaz (1987) pointed
out that thè specimens of this species, discovered in
Piedmont, could belong to a subspecies similar to P.
monspeliensis.
Portunites Bell, 1858, known to date from thè
Eocene of N Europe, N Africa, Japan and N America,
is reported for thè first time in Italy with P. rosen-
feldi. Schweitzer & Feldmann (2000) discussed Por¬
tunites Bell, 1858, known in thè fossil record with
eight species. Portunites rosenfeldi differs from thè
known fossil species in terms of thè location of thè
epibranchial rise creating a wide transverse convexity,
wider and angled at thè margins of thè protogastric
regions.
Finally, Titanocarcinus raulinianus A. Milne
Edwards, 1863, was reported by A. Milne Edwards
(1863), Lòrenthey (1898), and Lòrenthey & Beurlen
(1929) from thè middle Eocene of France and from
thè late Eocene of Hungary, while Ristori (1893), Di
Salvo (1933), and Beschin et al. (2000) reported this
species in Italy from thè lower Eocene of thè Gecche-
lina quarry of Monte di Malo (Vicenza), and from thè
middle Eocene of Monte Saraceno (Gargano, Puglia)
and Monreale (Sicily). Titanocarcinus includes many
species from thè Cretaceous to Miocene (Glaessner,
1969). W e pointed out that T. subovalis Ristori, 1896,
from thè Pliocene of Tuscany possessed a shape and
dorsal characters of thè carapace that are different
from those of Titanocarcinus, while T. sculptus Ri¬
stori, 1891, from thè Pliocene of Mucigliani (Siena), is
a synonym with Chlinocephalus demissifrons Ristori,
1886 (Garassino et al., 2004).
References
Allasinaz A., 1987 - Brachyura decapoda oligocenici
(Rupeliano) del Bacino Ligure Piemontese. Bol¬
lettino del Museo regionale di Scienze naturali di
Torino, Torino, 5 (2): 509-566.
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1988 - Raninidae del Terziario berico-lessineo
(Italia settentrionale). Lavori - Società Veneziana di
Scienze Naturali, Venezia, 13: 155-215.
Beschin C., Busulini A., De Angeli A., Tessier G. &
Ungaro S., 2000 - The fauna of thè Gecchelina
Quarry at Monte di Malo (Vicenza - Northern
Italy): a preliminary study. In: lst Workshop on
Mesozoic and Tertiary decapod crustaceans. Studi e
Ricerche - Associazione Amici del Museo - Museo
Civico “G. Zannato", Montecchio Maggiore: 7-12.
Collins J. S. H. & Dieni I., 1995 - New decapod crus¬
taceans from thè Cenomanian Rudist Limestones
of NE Italy. Bulletin of Mizunami Fossil Museum,
Mizunami, 22: 67-72.
Comaschi Caria I., 1950 - Crostacei Decapodi nel
Miocene (Elveziano) di Bosa in Sardegna. Ren¬
diconto del Seminario della Facoltà di Scienze
dell’Università di Cagliari, Cagliari, 20 (3-4):
324-327.
Comaschi Caria I., 1956 - I Crostacei miocenici
della Sardegna. Bollettino del Servizio Geologico
d’Italia, Roma, 78 (1-2): 283-290.
Dainelli G., 1915 - L’Eocene Friulano - Monografia
geologica e paleontologica. Editrici le “ Memorie
Geografiche” , Firenze.
De Angeli A. & Garassino A., 2006 - New reports of
decapod crustaceans from thè Mesozoic and Ceno-
zoic of Friuli-Venezia Giulia (NE Italy). Atti della
Società italiana di Scienze naturali e del Museo
civico di Storia naturale in Milano, Milano, 147
(2): 267-294.
Di Salvo G., 1933 - I Crostacei del Terziario inferiore
della provincia di Palermo. Giornale delle Scienze
Naturali ed Economiche di Palermo, Palermo, 37:
1-44.
Garassino A., De Angeli A., Gallo L. M. & Pasini G., 2004
- Brachyuran and anomuran fauna from thè Cenozoic
of Piedmont (NW Italy). Atti della Società italiana di
Scienze naturali e del Museo civico di Storia naturale
in Milano, Milano, 145 (2): 251-281.
Gemmellaro G. G., 1869 - Studi paleontologici sulla
fauna del calcare a Terebratula janitor del Nord
di Sicilia. Stabilimento Tipografico Lao, Palermo,
Parte I.
Glaessner M. F., 1969 - Crustacea Decapoda. In:
Treatise on Invertebrate Paleontology. Arthropoda
4 (2), Geologica! Society of America and University
of Kansas, Lawrence: R399-R533, R626-R628.
Lòrenthey I. E., 1898 - Beitràge zur Dekapodenfauna
des Ungarischen Tertiars. Termes-Fiizetek, Buda¬
pest, 21: 1-133.
Lòrenthey I. E., 1909 - Beitràge zur tertiàren Dekapo¬
denfauna Sardiniens. Mathematischen und Natur-
wissenschaftlichen Berichte aus Ungarn, Budapest,
24: 202-257.
Lòrenthey I. & Beurlen K., 1929 - Die fossilen Deka-
poden der Lànder Ungarischen Krone. Geologica
Hungarica, Serie Palaeontologica, Budapest, 3:
1-420.
Marras G. & Ventura G., 1991 - Crostacei decapodi
del Miocene di Sassari (Sardegna nord-occiden-
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
27
tale). Bollettino della Società Sarda di Scienze
Naturali, Sassari, 28: 105-119.
Milne Edwards A., 1862-1865 - Monographie des
Crustacés fossiles de la famille des Cancériens.
Annales de Science Naturelle, Zoologie, Paris, sér.
4, 18 (1862): 31-85; 20 (1863): 273-324; sér. 5, 1
(1864): 31-88; 3 (1865): 297-351.
Rasmussen H. W., 1971 - Echinoid and crustacean
burrows and their diagenetic significance in thè
Maastrichtian-Danian of Stevns Klint, Denmark.
Lethaia, Oslo, 4: 191-216.
Ristori G., 1888 - Alcuni Crostacei del Miocene
medio italiano. Atti della Società Toscana di Sci¬
enze Naturali, Pisa, 9: 212-219.
Ristori G., 1893 - Il Titanocarcinus raulinianus A.
M. Edw. negli strati nummulitici del Gargano. Atti
della Società Toscana di Scienze Naturali, Processi
verbali, Pisa, 7-8: 212-215.
Schweitzer C. E. & Feldmann R. M., 2000 - New fossil
portunids from Washington, USA, and Argentina,
and a re-evaluation of generic and family relation-
ships within thè Portunoidea Rafinesque, 1815
(Decapoda: Brachyura). Journal of Paleontology,
Lawrence, 74 (4): 636-653.
Swen K., Fraaije R. H. B., van der Zwaan G. J., 2001 -
Polymorphy and extinction of thè Late Cretaceous
burrowing shrimp Protocallianassa faujasi and
fìrst record of thè genera Corallianassa and Cal-
liax (Crustacea, Decapoda, Thalassinoidea) from
thè Cretaceous. Contributions to Zoology, The
Hague, 70 (2): 85-98.
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: agarassino@libero.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Rodney M. Feldmann, Robin Green & Carrie E. Schweitzer
An unusual Albian (Early Cretaceous) crab from Montana:
thè earliest eubrachyuran?
Discovery of a single specimen of a unique fossil
crab from thè Shell Creek Shale in Yellowstone County,
Montana, U.S.A., provides evidence that thè Eubrachyura
de Saint Laurent, 1980, may have appeared in thè fossil
record earlier than previously thought. The specimen
exhibits a plexus of anatomical features typical of both
primitive crabs within thè Podotremata Guinot, 1977,
and thè derived forms of thè Heterotremata Guinot, 1977,
embraced within thè Eubrachyura de Saint Laurent, 1980.
This combination of morphological features, which has
not been recognized previously, suggests that thè differ-
entiation of thè two major crab groups may have occurred
by a series of stages during thè latest Early Cretaceous
and thè early Late Cretaceous.
The specimen was collected along with a second, tiny
crab from a sequence of gray and black shale, thè Shell
Creek Shale, lying between sandy beds of thè Muddy
Sandstone and thè black, siliceous shales of thè Mowry
Shale (Eicher, 1 962), which is known from exposures in
centrai and eastem Montana and in Wyoming as well as
in thè subsurface of thè Bighom Basin. The Shell Creek
Shale has variously been considered a member of thè
Thermopolis Shale (eg. Moberly, 1960) or as a distinct
formation (Eicher, 1960, 1962; Condon, 2000). The Shell
Creek Shale was deposited in a deepening-upward basin
in which salinity increased stratigraphically upward
(Eicher, 1960). Twelve species of foraminiferans have
been identified from thè formation (Eicher, 1960); how-
ever, macrofossils are not common. Those that have been
identified include fish scales, turtle fragments, inoceramid
bivalves, and ammonites (Eicher, 1960, p. 27). Directly
relevant to this study, he also reported that Henry B. Rob-
erts, a prominent worker on fossil decapods, identified
two crab species, Homolopsis sp. and Dakoticancer sp.,
from localities in northem Bighom County, Wyoming.
Those specimens have not been examined yet to confimi
thè identification; however, one tiny crab was collected
in association with thè unusual crab under discussion.
This tiny crab may be a homolid and it will be important
to determine whether it is conspecific with thè specimen
Roberts referred to Homolopsis sp. Because all of thè
specimens, other than thè two crabs from Montana, are
from localities several miles to thè Southwest, it is dif-
fìcult to assess thè precise environmental setting, but it
can be inferred that thè crabs inhabited a normal marine
habitat on a fine-grained substratum.
The possible eubrachyuran specimen is preserved
as a carapace with stemum, maxillipeds, and mandibles
in living position. The abdomen and thoracic append-
ages are absent. It is preserved in a hard, dark matrix
and thè cuticle is altered to a dark color, reminiscent of
iron or manganese oxide. Although most of thè matrix is
extremely hard, some has oxidized and yields thè yellow-
orange streak of limonite. The dorsal surface of thè cara¬
pace is about 28 mm long and 32 mm wide, hexagonal in
outline, and weakly vaulted. Regions are moderately well
defined and some of them bear large, rimmed pits which
may have been thè sites of setal hair tufts. The stemum is
elongate ovoid, and complete sutures separate stemites 2-
3, 3-4, 5-6, 6-7 and 7-8. Stemites 4 and 5 are fused axially
and sutured laterally. The posteriormost somites, 7 and 8,
are reduced in width and length and are rotated dorsally.
Although not present, it appears that pereiopod 4 was
reduced in size, relative to thè preceeding ones, and that
pereiopod 5 was even smaller and was situated dorsally
or subdorsally. Superficially, thè outline of thè stemum
and thè degree to which stemites are fused is reminiscent
of some scyllarid slipper lobsters on one hand and of
thè dakoticancrid crabs on thè other. In either case, thè
generai conformation of thè stemum appears to be primi¬
tive. However, large orifices, interpreted to be vulvae, are
developed near thè axis of thè sixth abdominal somite - a
character typifying thè Heterotremata within thè Eubrach¬
yura as defined by de Saint Laurent (1980) and adopted by
Martin and Davis (2001). Thus, thè stemal architecture
does not conform to any previously known brachyuran,
leading to thè conclusion that thè specimen represents a
new family and genus of crab.
The combination of what are considered primitive
and derived characters exhibited by a single individuai
provides an opportunity for developing alternative evo-
lutionary scenarios. One interpretation would be that thè
definitional characters separating thè Podotremata from
thè Heterotremata arose in stages, rather than as a single
evolutionary event. There is, for example, no compelling
reason to argue that thè development of a vulva on thè 6th
stemite must of necessity have evolved simultaneously
with other changes in thè morphology of thè stemum and
thè conformation of thè 4th and 5th pereiopods defining thè
Heterotremata. This scenario would support thè conten-
tion that thè full array of derived characters arose over a
reasonably long period of time and that thè specimen at
hand simply documents one stage in that transition.
An alternative, and quite different, interpretation
would be that this organism represents a relatively early
experiment in thè evolution of brachyurans - one that did
not stand thè test of time. Schram (1980, 1983) discussed
thè concept of Bauplàne, body plans, in various arthropod
and non-arthropod groups. He postulated a morphologi¬
cal landscape based upon presence or absence of essen-
tial features, such as uniramous or biramous thoracic
appendages, presence or absence of a brood pouch, etc.
When arrayed as a morphological landscape, it became
clear that several of thè Bauplàne embraced recognizable
31® SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
29
clades, some morphological combinations have never
been recognized, and some combinations emerged early
in thè radiation of thè clade and were apparently unsuc-
cessful. Applying this scenario, thè Albian crab from
Montana would have represented an early and failed
experiment in thè radiation of thè Brachyura. Testing
these two hypotheses must involve discovery, collection,
and study of additional material.
References
Condon S. M., 2000 - Stratigraphic framework of Lower
and Upper Cretaceous rocks in centrai an eastem
Montana. U. S. Geological Survey Digital Data
Series, Washington, DDS57.
Eicher D. L., 1960 - Stratigraphy and micropaleontol-
ogy of thè Thermopolis Shale. Peabody Museum of
Naturai History, Yale University, Bulletin, New Haven
Connecticut, 15: 1-126.
Eicher D. L., 1962 - Biostratigraphy of thè Thermopolis,
Muddy, and Shell Creek formations. Wyoming Geo¬
logical Association Guidebook, Laramie, Wyoming,
17* Annual Field Conference: 72-93.
Guinot D., 1977 - Propositions pour une nouvelle classifì-
cation des Crustacés Décapodes Brachyoures. Comp-
tes Rendus des Seances de l’Academie des Sciences,
Series 3, Paris, 285: 1049-1052.
Haan W. de, 1833-1850 - Crustacea. In: Fauna Japonica
sive Description Animalium, quae in Itinere per Japo-
niam, Jussu et Auspiciis Superiorum, qui Summum in
India Batava Imperium Tenent, Suscepto, Annis 1823-
1830 Collegit, Notis, Observationibus et Adumbratio-
nibus Illustravit. P. F. von Siebold & J. Miiller (eds.).
Lugduni-Batavorum (=Leiden).
Martin J. W. & Davis G. E., 2001 - An updated classifica-
tion of thè Recent Crustacea. Naturai History Museum
of Los Angeles County, Science Series, Los Angeles,
39: 1-124.
Moberly R. Jr., 1960 - Morrison, Cloverly, and Sykes
Mountain formation, northem Bighom Basin, Wyo¬
ming and Montana. Geological Society of America
Bulletin, Boulder, Colorado, 71: 1137-1176.
Saint Laurent M. de, 1980 - Sur la classification et la
phylogénie des Crustacés Décapodes Brachyoures.
I. Podotremata Guinot, 1977, et Eubrachyura sect.
nov. Comptes Rendus Hebdomadaries des seances de
l’Academie des Sciences Sèrie D, Paris, 290: 1265-
1268.
Schram F. R., 1980 - On thè Classification of Eumala-
costraca. Journal of Crustacean Biology, Lawrence,
Kansas, 1: 1-10.
Schram F. R., 1983 - Method and madness in phylogeny.
In: Crustacean Phylogeny. F. R. Schram (ed.). A. A.
Balkema, Rotterdam: 331-350.
Rodney M. Feldmann - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rfeldman@kent.edu
Robin Green - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rgreen8@kent.edu
Carrie E. Schweitzer - Dept. of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, U.S.A.
e-mail: cschweit@kent.edu
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Rodney M. Feldmann& Carrie E. Schweitzer
A compilation of Decapoda collections in Museums of thè world
Museum collections offer an often underappreciated
and untapped source of material that can yield a wealth
of information on phylogeny, systematics, paleoecology,
paleobiogeography, and evolution (Allmon, 2005). It has
been estimated that there is more unpublished material
housed in museums than has been described and that this
may seriously hinder phylogenetic and statistical analyses
of thè fossil record (Teichert et al., 1987). Some of thè
material, especially in older collections, may have been
collected from localities that are no longer accessible or
that simply no longer exist (i.e., many 19th century locali¬
ties). Museum collections are unparalleled sources of
comparative material, and studying type material is thè
single best means by which to conduct systematic and
phylogenetic studies. Large paleoecological studies are
often most efftciently conducted using large museum col¬
lections of specimens that were amassed over many years
from localities that may yield few specimens on any given
day. Thus, it is cruciai to evaluate thè existing collections
of fossil decapod crustaceans and to be cognizant of thè
location of important type, geographic, and stratigraphic
collections.
Locating existing type and other material in museums
can be a diftkult task. Many museums do not have pub-
lished catalogs, or if so, they are not on line for access
to anyone, anywhere in thè world. The location of many
important collections of fossil decapod Crustacea has long
been unknown, such as most of thè material of Hermann
von Meyer and Victor Van Straelen. In an effort to locate
this and other decapod material, we constructed a ques-
tionnaire and sent it to a large number of museums, large
and small, that we believed might house collections of
fossil decapods. To date, thè response has been excellent.
We have received responses from 30 institutions. In addi-
tion, we compiled information from published catalogues
or our own personal research or communication from
nine museums for a total of 39 museums in 14 countries
on four continents. Unfortunately, most of thè von Meyer
material has yet to be located, but we have located many
collections that have been little known.
Our questionnaire sought thè following information:
name of museum or institution, curator of decapod or
invertebrate collection, address, estimate of number of
catalogued specimens of decapods, estimate of number of
type specimens of decapods, collections of notable deca¬
pod workers, decapod collections from specific areas,
ages, or formations, and whether a decapod catalogue
was available. We also asked for leads on other museum
collections to query. Thus, we continue to seek informa¬
tion on decapod collections, and following, we report thè
information we have collected thus far. We include some
museums for which we received no information but for
which a published catalogue exists or for which we pos-
sessed information based upon our own research or per¬
sonal communication. Museums for which information is
derived solely from thè published catalogue are marked
with an asterisk (*), and those for which thè information
was derived from our own work or personal contacts are
marked with a doublé asterisk (**).
Europe
Austria
Museum: Naturhistorisches Museum in Wien, Geolo-
gisch-Palaontologische Abteilung
Curator: Dr. Ortwin Schultz
Address: Burgring 7, 1010 Wien, Austria
Catalogue available?: no
Estimate of number of catalogued decapods: many
Estimate of number of type [and figured] specimens of
decapods: many
Collections of notable workers: F. Bachmayer, F. Blas-
chke, M. F. Glaessner
Collections from a specific area, age, or formation:
Jurassic of Austria, many other European localities
France
Museum: Museum national d’Histoire naturelle (Paris)**
Curator: Dr. Jean Paul Saint Martin
Address: Paris 5ème, Jardin des Plantes 2, rue Buffon,
Paris, France
Catalogue available?: no, but see Secretan (1964) for
Madagascar material
Estimate of number of catalogued decapods: many
Estimate of number of type [and figured] specimens of
decapods: many
Collections of notable workers: A. Milne Edwards, S.
Secretan, J. M. Remy
Collections from a specific area, age, or formation: France;
Jurassic and Cretaceous of Madagascar; Africa
Museum: Musée Vert, Musée d’Histoire naturelle du
Mans
Curator: Dr. Nicolas Morel
Address: 204, avenue Jean-Jaurès 72000, Fe Mans,
France
Catalogue available?: no, but see A. Milne Edwards &
Brocchi (1879) and Van Straelen (1936)
Estimate of number of catalogued decapods: at least 20
Estimate of number of type [and figured] specimens of
decapods: unknown
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
31
Collections of notable workers: a large portion of thè
collection was lent to thè Musée Royal d’Histoire
Naturelle de Belgique and was subsequently studied
and published by Van Straelen (1936). This material
has apparently been lost (N. Morel, personal com-
munication)
Collections from a specific area, age, or formation: Ceno-
manian stratotype of Le Mans, France
Museum: Observatoire des Sciences de TUnivers de Gre¬
noble (OSUG), Université Joseph Fourier
Curator: Dr. Emmanuel Robert
Address: Institut Dolomieu, 15, rue Maurice Gignoux,
38031 Grenoble cedex, France
Catalogue available?: yes
Estimate of number of catalogued decapods: about 32
Estimate of number of type [and figured] specimens of
decapods: 3 holotypes
Collections of notable workers: V. Van Straelen
Collections from a specific area, age, or formation:
Jurassic and Cretaceous of France
Museum: Museum d’Histoire Naturelle de Nantes
Curator: Dr. Serge Regnault
Address: MHNN 12, rue Voltaire, 44000 Nantes, France
Catalogue available?: no
Estimate of number of catalogued decapods: none given
Estimate of number of type [and figured] specimens of
decapods: 1 holotype
Collections of notable workers: none given
Collections from a specific area, age, or formation: none
given
Museum: Collections de Géologie, Université Claude
Bernard Lyon
Curator: Dr. Abel Prieur (Directeur des Collections de
Géologie)
Address: Université Claude Bernard Lyon, CNRS UMR
5125 PEPS, Paléoenvironments et Paléobiosphère,
Bàtiment Géode, 2, rue Dubois, 69622 Villeurbanne
Cedex, France
Catalogue available?: no
Estimate of number of catalogued decapods: about 6
Estimate of number of type [and figured] specimens of
decapods: 1 holotype, 1 paratype
Collections of notable workers: J. P. Saint Martin & P.
Muller (1988)
Collections from a specific area, age, or formation:
Miocene of Algeria
Museum: Collection de l’Ecole des Mines de Paris depose
à Lyon
Catalogue available?: no
Estimate of number of catalogued decapods: about 4
Estimate of number of type [and figured] specimens of
decapods: 3 holotypes, 1 paratype
Collections of notable workers: V. Van Straelen, A.
Oppel
Collections from a specific area, age, or formation:
Jurassic and Cretaceous of France
Museum: Service de Valorisation des Collections de Géo¬
logie (SERVACO-Géol), Université de Bourgogne
Curator: Dr. Jéròme Thomas
Address: SERVACO-Géol, UMR uB-CNRS 5561 Bio-
géosciences, UFR des Sciences de la Terre, 6, boule¬
vard Gabriel, 21000 Dijon, France
Catalogue available?: http://transtyfipal.u-bourgogne.fr
Estimate of number of catalogued decapods: unknown
Estimate of number of type [and figured] specimens of
decapods: 1 in University of Burgundy
Collections of notable workers: none
Collections from a specific area, age, or formation: none
Germany
Museum: Bavarian State Collection for Palaeontology
and Geology (Bayerische Staatsammlung fur Palàon-
tologie und historische Geologie)
Curator: Dr. Martin Nose
Address: Richard- Wagner-StraBe 10, D-80333 Munchen,
Germany
Catalogue available: not yet
Estimate of number of catalogued decapods: 1,500-2,000
(preliminary)
Estimate of number of type [and figured] specimens of
decapods: 300-500 (preliminary)
Collections of notable workers: R. Forster, W. Moericke,
G. G. zu Munster, C. A. Oppel, G. Wehner
Collections from a specific area, age, or formation: Upper
Jurassic of Solnhofen and Eichstàtt
Museum: Forschungsinstitut Senckenberg
Curator: Dr. Alan Lord and Claudia Franz (Technical
Assistant)
Address: Sektion Mikropalaeontologie I, Forschungsin¬
stitut Senckenberg, Senckenberganlage 25, 60325
Frankfurt am Main, Germany
Catalogue available?: yes, but not yet complete
Estimate of number of catalogued decapods: about 1,000
specimens
Estimate of number of type [and figured] specimens of
decapods: 1 holotype and published material of M.
Glaessner, H. Malz, O. Kuhn, K. A. Frickinger, V. Van
Straelen, H. von Meyer
Collections of notable workers: H. von Meyer, H. Malz
Collections from a specific area, age, or formation:
Jurassic (Solnhofen-type limestones) of Germany
Museum: Museum fur Naturkunde Berlin, Palàon-
tologisches Museum, Humboldt-Universitàt zu
Berlin**
Curator: Dr. Christian Neumann
Address: Museum fur Naturkunde, der Humboldt-Uni¬
versitàt zu Berlin, Invalidenstrasse 43, D- 10115
Berlin, Germany
Catalogue available?: no
Estimate of number of catalogued decapods: unknown,
at least 100
Estimate of number of type [and figured] specimens
of decapods: 51 holotypes and figured specimens
(recorded by RMF and CES, June, 2001)
Collections of notable workers: A. Bittner, A. Desmarest,
O. Fraas, H. von Meyer, G. G. zu Munster, F. Noetling,
E. F. von Schlotheim
Collections from a specific area, age, or formation:
Jurassic (Solnhofen-type limestones) of Germany;
Eocene of Italy (Bittner)
32
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Museum: Staatliches Museum fur Naturkunde Stuttgart
Curatori Dr. Giinter Schweigert (Triassic), Dr. Gert Dietl
(Jurassic), Dr. Michael W. Rasser (Cenozoic)
Address: Staatliches Museum fur Naturkunde, Rosenstein
1, 70191 Stuttgart, Germany
Catalogue available: no
Estimate of number of catalogued decapods: unknown
Estimate of number of type [and figured] specimens of
decapods: about 40 holotypes
Collections of notable workers: O. Fraas collection; R.
Mundlos collection
Collections from a specific area, age, or formation:
Triassic and Jurassic of SW Germany, especially
lithographic limestones of Solnhofen and Nusplingen;
Eocene of northem Germany (Helmstedt) and Egypt;
Cretaceous of Lebanon
Hungary
Museum: Fòldtani Intézet (Geological Survey)**
Curatori Dr. Pài Miiller
Address: Stefania ùt 14, H-1143 Budapest, Hungary
Catalogue available: contact Dr. Pài Miiller
Estimate of number of catalogued decapods: 370
Estimate of number of type [and figured] specimens of
decapods: 33 holotypes
Collections of notable workers: E. Lòrenthey
Collections from a specific area, age, or formation: Ceno¬
zoic of Hungary
Museum: Természettudomànyi Muzeum Fòldés Òslénytàr
(Naturai History Museum of Hungary)**
Curatori Dr. Pài Miiller
Address: contact Dr. Pài Miiller at Stefània ùt 14 H-1143
Budapest Hungary
Catalogue available?: contact Pài Miiller
Estimate of number of catalogued decapods: 560 cata¬
logued Miocene, about 800 Eocene
Estimate of number of type [and figured] specimens of
decapods: at least 11 Miocene holotypes, about 88
Eocene holotypes
Estimate of number of uncatalogued decapods: types
and other material from Miiller (1974) are not yet
catalogued and are in two separate boxes in a large,
low drawer.
Collections of notable workers: P. Miiller
Collections from a specific area, age, or formation: Ceno¬
zoic of Hungary
Italy
Museum: Museo Civico di Storia Naturale di Milano
Curatori Dr. Alessandro Garassino, Dr. Giorgio Teruz-
zi
Address: Corso Venezia, 55, 20121 Milano, Italy
Catalogue available?: see De Angeli & Garassino
(2006)
Estimate of number of catalogued decapods: 2,680
Estimate of number of type [and figured] specimens of
decapods: 49 holotypes, 125 patypes and 187 figured
Collections of notable workers: A. Garassino, G. Te-
ruzzi
Collections from a specific area, age, or formation: Meso-
zoic of Italy, Lebanon, and Madagascar
Museum: Museo Civico “G. Zannato” di Montecchio
Maggiore (Vicenza)
Curatori Dr. Viviana Frisone
Address: P. le Marconi 15, 1-36075 Montecchio Maggiore
(Vicenza), Italy
Catalogue available: yes
Estimate of number of catalogued decapods: 1,450
Estimate of number of type [and figured] specimens of
decapods: about 63 holotypes, about 242 paratypes
Collections of notable workers: C. Beschin, A. Busulini,
A. De Angeli, G. Tessier
Collections from a specific area, age, or formation: Ceno¬
zoic of Italy
Museum: Museo Civico D. Dal Lago
Curatori Dr. Bemardetta Pallozzi, Dr. Dario Savi
Address: Palazzo Festari, Corso Italia 63, 36078 Valda-
gno, (Vicenza), Italy
Catalogue available?: yes
Estimate of number of catalogued decapods: 16
Estimate of number of type [and figured] specimens of
decapods: 2 paratypes
Collections of notable workers: none
Collections from a specific area, age, or formation:
middle Eocene of thè localities Grola Cave and Monte
di Malo, near Vicenza, Italy
Museum: Museo del Dipartimento per lo Studio del Ter¬
ritorio e delle sue Risorse, Università di Genova
Curatori Dr. Maria Cristina Bonci
Address: Dip.Te.Ris., Università di Genova, Corso
Europa 26, 16132 Genova, Italy
Catalogue available?: see De Angeli & Garassino (2006)
Estimate of number of catalogued decapods: 5
Estimate of number of type [and figured] specimens of
decapods: 3 holotypes
Collections of notable workers: none
Collections from a specific area, age, or formation: Fer¬
rando Collections - Sassello (Savona, Liguria, Italia
settentrionale) - Oligocene - Tertiary Piedmont Basin,
Molare Formation
Museum: Museo di Storia Naturale dell’Università, degli
Studi di Firenze, Sezione di Geologia e Paleontologia
Curatori Dr. Laura Delle Cave
Address: Museo di Storia Naturale dell’Università, degli
Studi di Firenze, Sezione di Geologia e Paleontologia,
Via G. La Pira, 4, 50121 Firenze, Italy
Catalogue available?: see Delle Cave (1981)
Estimate of number of catalogued decapods: unknown
Estimate of number of type [and figured] specimens of
decapods: 22 holotypes, some of which may be lost
Collections of notable workers: G. Checchia-Rispoli, G.
Ristori
Collections from a specific area, age, or formation: Neo¬
gene of Italy
Museum: Museo del Dipartimento di Geologia, Paleonto¬
logia e Geofisica dell’Università di Padova
Curatori Dr. Mariagabriella Fomasiero
Address: Museo del Dipartimento di Geologia, Paleon¬
tologia e Geofìsica dell’Università di Padova, Via
Giotto, 1-35137 Padova, Italy
Catalogue available?: yes
3“ SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
33
Estimate of number of catalogued decapods: 240
Estimate of number of type [and figured] specimens of
decapods: 1 1 holotypes, 4 figured
Collections of notable workers: R. Fabiani, S. Secretan
Collections from a specific area, age, or formation:
Eocene of Italy
The Netherlands
Museum: Oertijdmuseum De Groene Poort
Curatori Dr. René H. B. Fraaije
Address: Bosscheweg 80, 5283 WB Boxtei, The Nether¬
lands
Catalogue available?: not yet
Estimate of number of catalogued decapods: 2,500
Estimate of number of type [and figured] specimens of
decapods: 30 holotypes
Collections of notable workers: R. H. B Fraaije, B. van
Bakel
Collections from a specific area, age, or formation: type
Maastrichtian, Cenozoic of western Europe, Late
Jurassic of southern Germany and UK
Poland
Museum: Institute of Paleobiology, Polish Academy of
Sciences
Curatori Dr. Maria Aleksandra Bitner
Address: Institute of Paleobiology, ul. Twarda 51/55, 00-
818, Warszawa, Poland
Catalogue available?: yes
Estimate of number of catalogued decapods: about 300
and about 100 uncatalogued specimens
Estimate of number of type [and figured] specimens of
decapods: 2 holotypes
Collections of notable workers: A. Gazdzicki
Collections from a specific area, age, or formation:
Miocene Cape Melville Formation, King George
Island, Antarctica; Eocene La Meseta Formation,
Seymour Island, Antarctica; Upper Jurassic of
Poland
Switzerland
Museum: Musée Cantonal de Géologie, Lausanne
Curatori Dr. Robin Marchant
Address: UNIL-Anthropole, CH-1015 Lausanne, Swit¬
zerland
Catalogue available?: see Septfontaine (1995). PDF
available at www.unil.ch/webdav/site/mcg/shared/
Types_Paleonto_MGL.pdf
Estimate of number of catalogued decapods: 50
Estimate of number of type [and figured] specimens of
decapods: 8 holotypes
Collections of notable workers: M. de Tribolet
Collections from a specific area, age, or formation: Swiss
Jurassic
Museum: Muséum d’Histoire Naturelle de la Ville de
Genève
Curatori Dr. Lionel Cavin
Address: Département de Géologie et Paléontologie,
Muséum d’Histoire Naturelle de la Ville de Genève,
CP 6434 1211 Geneve 6, Switzerland
Catalogue available?: in part, A. Favre collection is pub-
lished (Benier & Berset, 1989)
Estimate of number of catalogued decapods: 50
Estimate of number of type [and figured] specimens of
decapods: 1 figured
Collections of notable workers: V. Van Straelen
Collections from a specific area, age, or formation: Creta-
ceous of France and UK, Jurassic of Germany
United Kingdom
Museum: National Museums of Scotland, Edinburgh
Curatori Dr. Lyall I. Anderson
Address: Department of Naturai Sciences, National
Museums of Scotland, Building 8, West Granton
Centre, 242, West Granton Road, Edinburgh, EH5 1 JA
Scotland, UK
Catalogue available?: yes but not available over thè inter¬
net yet
Estimate of number of catalogued decapods: 98
Estimate of number of type [and figured] specimens of
decapods: none
Collections of notable workers: A. Copland Hutchison
Collections from a specific area, age, or formation:
Solnhofen (Jurassic) of Germany; Gault Clay (Cre-
taceous) of Folkestone, Kent, England; London Clay
(Eocene) of London, England and thè Isle of Sheppey
Museum: The Naturai History Museum (London)
Curatori Dr. Andrew Ross
Address: Department of Palaeontology, The Naturai His¬
tory Museum, Cromwell Road, S. Kensington, SW7
5BD London, UK
Catalogue available?: see Morris (1980)
Estimate of number of catalogued decapods: 6,000
Estimate of number of type [and figured] specimens of
decapods: 500 holotypes
Collections of notable workers: T. Bell, J. S. H. Collins,
S. F. Morris, T. H. Withers, H. Woods, H. Woodward,
C. W. Wright
Collections from a specific area, age, or formation:
Jurassic, Cretaceous, and Eocene of UK; many former
British colonies and protectorates
Museum: Sedgwick Museum of Earth Sciences
Curatori Dr. Liz Harper (Honoree Curator), Dan Pember-
ton (Collections Manager), Mathew Lowe (Collec¬
tions Assistant)
Address: Sedgwick Museum of Earth Sciences, Univer¬
sity of Cambridge, Downing Street, CB2 3EQ Cam¬
bridge, UK
Catalogue available?: yes
Estimate of number of catalogued decapods: 1 ,2 1 8
Estimate of number of type [and figured] specimens of
decapods: 54 syntypes, 19 holotypes, 4 metatypes,
150 figured
Collections of notable workers: T. Bell, Carter, G. Man-
tell, C. W. Wright
Collections from a specific area, age, or formation: Creta¬
ceous and Cenozoic of Britain
Museum: University Museum of Naturai History (Oxford)
Curator: Dr. Erik Seiffert
Address: Parks Road, OX1 3PW Oxford, UK
34
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Catalogue available?: http://www.oum.ox.ac.uk/collect/
geology2.htm
Estimate of number of catalogued decapods: about 1,000
Estimate of number of type [and figured] specimens of
decapods: 35 holotypes
Collections of notable workers: T. Bell, J. S. H. Collins, J.
Phillips, W. J. and S. Quayle
Collections from a specific area, age, or formation:
Jurassic, Cretaceous, and Eocene of UK; Cretaceous
of France
Asia
Japan
Museum: Mizunami Fossil Museum
Curator: Dr. Hiroaki Karasawa
Address: Yamanouchi, Akeyo, Mizunami, Gifu 509-6132,
Japan
Catalogue available?: http://www2.city.mizunami.gifii.jp/
mizunami/dbtope . html
Estimate of number of catalogued decapods: 633
Estimate of number of type [and figured] specimens of
decapods: 76 holotypes, 91 paratypes
Collections of notable workers: H. Karasawa
Collections from a specific area, age, or formation: all
areas of Japan
Museum: Naturai History Museum and Institute, Chiba
Curator: Dr. Hisayoshi Kato
Address: 955-2 Aoba-cho, Chuo-ku, Chiba 260-8682, Japan
Catalogue available?: yes
Estimate of number of catalogued decapods: 1,500
Estimate of number of type [and figured] specimens of
decapods: 1 holotype, 17 paratypes
Collections of notable workers: H. Kato
Collections from a specific area, age, or formation: north-
eastem Japan
Synopsis of other Japanese Museums (from H. Kato):
Tohoku University Museum, Institute of Geology and
Paleontology, Sendai: 800 numbered specimens, of
which about 50 are holotypes or figured specimens of
R. Imaizumi or H. Kato
Hokkaido University Museum: 20 holotypes and figured
specimens of T. Nagao
National Science Museum: about 40 specimens of which
6 are holotypes or figured specimens of M. Takeda and
I. Fujiyama
Saitama Museum of Naturai History: 200 specimens from
Miocene Chichibu Basin
Gunma Museum of Naturai History: about 50 specimens
including 2 patypes, 23 figured
Kanagawa Museum of Naturai History: about 200 speci¬
mens including 1 holotype, 1 paratype, 23 figured
North America
Mexico
Museum: Colecion Nacional de Paleontologia, Instituto
de Geologia, UNAM
Curator: Dr. Maria del Carmen Perrilliat
Address: Instituto de Geologia, UNAM, Ciudad Universi¬
taria, Coyoacan, Mexico DF 04510, Mexico
Catalogue available?: yes
Estimate of number of catalogued decapods: 150
Estimate of number of type [and figured] specimens of
decapods: 10 holotypes, 30 paratypes
Collections of notable workers: F. J. Vega
Collections from a specific area, age, or formation: Creta¬
ceous and Paleocene of northeastem Mexico
United States of America
Museum: Camegie Museum of Naturai History**
Curator: Albert Kollar (Collections Manager)
Address: Section of Invertebrate Paleontology, Camegie
Museum of Naturai History, 4400 Forbes Avenue,
Pittsburgh, Pennsylvania 15213 USA
Catalogue available?: yes
Estimate of number of catalogued decapods: about 600
Estimate of number of type [and figured] specimens of
decapods: 1 1 holotypes, 23 paratypes
Collections of notable workers: R. M. Feldmann, C. E.
Schweitzer, F. J. Vega
Collections from a specific area, age, or formation: Juras¬
sic Solnhofen-type limestones of Germany; Eocene
of Britain; Eocene Castle Hayne Formation of north
Carolina, USA; Cretaceous of Western Interior north
America; Paleogene of Washington, USA; Cretaceous
and Eocene of Latin America
Museum: The Field Museum
Curator: Dr. Scott Lidgard (Associate Curator, Fossil
Invertebrates), Dr. Peter Wagner (Associate Curator,
Fossil Invertebrates)
Address: Department of Geology, The Field Museum,
1400 S. Lake Shore Drive, Chicago, IL 60605 USA
Catalogue available?: no
Estimate of number of catalogued decapods: 466 lots
Estimate of number of type [and figured] specimens of
decapods: 254 lots
Collections of notable workers: none
Collections from a specific area, age, or formation: Juras¬
sic Solnhofen of Germany, Eocene Green Ri ver For¬
mation of western USA
Museum: Museum of Comparative Zoology, Harvard
University*
Curator: Dr. Fred Collier
Address: Museum of Comparative Zoology, Harvard
University, 26 Oxford Street, Cambridge, MA 02138-
2902, USA
Catalogue available?: see Rolfe (1963)
Estimate of number of catalogued decapods: about 39
Estimate of number of type [and figured] specimens of
decapods: 2 holotypes, 1 paratype, 3 syntypes, 3 fig¬
ured
Collections of notable workers: W. Stimpson, A. F. Eser
Collections from a specific area, age, or formation: Ger¬
many (see Rolfe, 1963)
Museum: Peabody Museum of Naturai History*
Curator: Dr. Susan Butts (Senior Collections Manager)
Address: Yale University, New Haven, CT 06520-8118, USA
3"° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
35
Catalogue available?: see White (1996)
Estimate of number of catalogued decapods: about 119
Estimate of number of type [and figured] specimens of
decapods: 3 holotypes, 1 paratype
Collections of notable workers: M. J. Rathbun, H. B.
Roberts
Collections from a specifìc area, age, or formation: Cre-
taceous and Cenozoic of Pacific north America; Cre-
taceous of east Coastal USA; Jurassic Solnhofen-type
limestones of Germany
Museum: San Diego Naturai History Museum
Curatori Dr. Scott Rugh (Collections Manager Inverte¬
brate Fossils)
Address: P. O. Box 121390, San Diego, California 92112-
1390 USA
Catalogue available?: yes
Estimate of number of catalogued decapods: 1,213
records representing 1 0,466 specimens
Estimate of number of type [and figured] specimens of
decapods: 85 holotypes
Collections of notable workers: G. A. Bishop, M, J. Rath¬
bun, C. E. Schweitzer, and R. M. Feldmann
Collections from a specifìc area, age, or formation: Creta-
ceous and Cenozoic of Pacific north America
Museum: United States National Museum of Naturai His¬
tory, Smithsonian Institution**
Curatori Warren C. Blow (Museum Specialist)
Address: Department of Paleobiology, National Museum
of Naturai History, P. O. Box 37012, NHB, MRC121,
Washington, D.C. 20013-7012 USA
Catalogue available?: yes
Estimate of number of catalogued decapods: unknown
Estimate of number of type [and figured] specimens of
decapods: many
Collections of notable workers: G. A. Bishop, W. C.
Blow, R. M. Feldmann, M. J. Rathbun, C. E. Sch¬
weitzer, H. B. Stenzel, A. B. Tucker
Collections from a specifìc area, age, or formation: north
America, centrai America, Cretaceous and Cenozoic
of Pacific north America
Oceania
Australia
Museum: Museum Victoria
Curatori Dr. David Holloway
Address: GPO Box 666, Melbourne, Victoria 3001, Aus¬
tralia
Catalogue available?: printed catalogue
Estimate of number of catalogued decapods: 340
Estimate of number of type [and figured] specimens of
decapods: 8 holotypes, 14 figured
Collections of notable workers: R. J. F. Jenkins, F. A.
Cudmore
Collections from a specifìc area, age, or formation: Ceno¬
zoic of southern Australia; Jurassic Solnhofen of Ger¬
many; Cretaceous of southern UK
New Zealand
Museum: Paleontology Collection, University of Auck¬
land
Curatori Dr. Neville Hudson
Address: School of Geography, Geology and Environ-
mental Science (SGGES), University of Auckland,
Private Bag 92019, Auckland Mail Centre, Auckland
1 142, New Zealand
Catalogue available?: yes
Estimate of number of catalogued decapods: 250-500
Estimate of number of type [and figured] specimens of
decapods: 2 holotypes
Collections of notable workers: none
Collections from a specifìc area, age, or formation: Paleo¬
gene and Neogene of New Zealand
References
Allmon W. D., 2005 - The importance of museum col¬
lections in paleobiology. Paleobiology , Lawrence,
Kansas, 31: 1-5.
De Angeli A. & Garassino A., 2006 - Catalog and Bibli-
ography of thè fossil Stomatopoda and Decapoda from
Italy. Memorie della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale di Milano ,
Milano, 35 (1): 1-95.
Benier C. & Berset S., 1989 - Les collections du Departe-
ment de Géologie et de Paléontologie des Invertebres
du Museum d’Histoire Naturelle de Genève. 35. La
Collection A. Favre (Cephalopoda, Trilobita, Deca¬
poda, Cirripedia). Revue de Paléobiologie, Geneva,
voi. spec. no. 3: 115-142.
Delle Cave L., 1981 - Catalogue of type specimens in
thè Invertebrate Palaeontological Collections of thè
Museum of Geology and Palaeontology of thè Uni¬
versity of Florence. (Italy). Crustacea, Decapoda. Atti
della Società Toscana di Scienze Naturali, Memorie,
Pisa, Serie A, 88: 43-50.
Milne Edwards A. & Brocchi P., 1879 - Note sur quel-
ques Crustacés fossils appartenant au groupe des
Macrophthalmiens. Bulletin de la Société Philomathi-
que de Paris, Paris, 7 (3): 113-117.
Morris S. F., 1980 - Catalogue of thè Type and Figured
specimens of fossil Crustacea (excl. Ostracoda), Che¬
licerata, Myriapoda and Pycnogonida in thè British
Museum (Naturai History). British Museum {Naturai
History), London.
Miiller R, 1974 - Decapoda (Crustacea) fauna a budapesti
miocénbòl (2). Foldtani Kòzlòny, Budapest, 104 (1):
275-287.
Rolfe W. D. I., 1963 - Catalogue of type specimens in thè
invertebrate paleontological collections of thè Museum
of Comparative Zoology. Arthropoda. Bulletin of thè
Museum of Comparative Zoology at Harvard College,
Cambridge, Massachusetts, 129 (7): 369-398.
Saint Martin J.-P. & Miiller R, 1988 - Les Crustacés
Décapodes du Messinien récifal d’Oranie (Algérie).
Geobios, Lyon, 21 (2): 251-257.
36
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Secretati S., 1964 - Les Crustacés Décapodes du Jurassi-
que supérieur et du Crétacé de Madagascar. Mémoires
du Muséum National d’Histoire Naturelle , Sèrie C,
Sciences de la Terre, Paris, 14: 1-223.
Septfontaine M., 1995 - Catalogue des types paléontolo-
giques deposes au Musée cantonal de Géologie, Lau¬
sanne. Mémoires de Géologie , Lausanne, 26: 1-78.
Teichert C., Sweet W. C & Boucot A. J., 1987 - The
unpublished fossil record: implications. Senckenber-
giana Lethaea, Frankfurt am Main, 68 (1/4): 1-19.
Van Straelen V., 1936 - Crustacés Décapodes nouveaux
ou peu connus de Fépoque Crétacique. Musée Royal
d’Histoire Naturelle de Belgique, Brussels, 12 (45):
1-40.
White R. D., 1996 - A type catalog of fossil invertebrates
(Arthropoda: Crustacea) in thè Yale Peabody Museum.
Postilla, New Haven, Connecticut, 210: 1-19.
Rodney M. Feldmann - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rfeldman@kent.edu
Carrie E. Schweitzer - Dept. of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, U.S.A.
e-mail: cschweit@kent.edu
3"1 Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
René H. B. Fraaije, Barry W. M. Van Bakel & John W. M. Jagt
A new species of Goniocypoda and thè first record of
Glyphithyreus wetherelli (Bell, 1858) (Decapoda, Brachyura)
from thè Eocene of Nieuwvliet-Bad, The Netherlands
The beaches at Nieuwvliet-Bad and Cadzand in thè
province of Zeeland (The Netherlands) and at Knokke
(northwest Belgium) are renowned for thè Cenozoic
fossil assemblages (mostly vertebrate and molluscan
remains, but also echinoderms and crabs) that are
washed ashore there. Just offshore, strata of late Eocene,
early-middle Oligocene and late Miocene age crop out,
and through periodic storms and daily tides fossils from
these levels are eroded loose and washed ashore. The
result is a near-constant supply of new material, in par-
ticular numerous shark teeth and molluscan shells (Lin-
demann & Fraaije, 2003). During thè 1980s and 1990s,
deeper-lying Cenozoic sands were used as supplemen-
tary material for beach protection in this area, notably at
Knokke and Cadzand. This resulted in an admixture of
fossils from those sands and from autochthonous strata
offshore. Phosphatic internai moulds of decapod crus-
taceans, mostly crabs, are not uncommon in this beach
material. Until now, only two species of crab, Coeloma
(C.) balticum and Harpactocarcinus sp. (= Eocarpilius
sp., as interpreted herein) were recorded (Fraaye, 1998;
Fraaije, 2003) from those localities, but these occur
in comparatively large numbers, and are sought-after
objects for collectors. The present record of a new spe¬
cies of Goniocypoda adds to thè list of post-Paleocene
decapod crustaceans currently known from Dutch terri-
tory (Tab. 1).
Systematic palaeontology
For higher-level classification of decapod crustaceans,
we follow Martin & Davis (2001).
Superfamily Xanthoidea MacFeay, 1838
Family Hexapodidae Miers, 1886
Genus Goniocypoda H. Woodward, 1867
Type species: Goniocypoda edwardsi H. Woodward,
1 867, by monotypy.
Stratigraphic range: ?Fatest Cretaceous (Maastrich-
tian); Paleocene-Eocene (see Schweitzer & Feldmann,
2001; Schweitzer, 2005; Guinot, 2006).
Goniocypoda verheyeni n. sp.
Diagnosis: carapace small, subrectangular in outline,
much wider than long; carapace regions poorly marked;
very wide orbitofrontal margin; posterior margin nearly
straight; large hook-shaped epimeral muscle scars; two
nodes at base of rostrum.
Derivation of name: in honour of Ivo Verheyen
(Moergestel, thè Netherlands), who collected and donated
thè type, and sole specimen known to date.
Material: thè holotype, a phosphatic internai mould,
is MAB k.2440, in thè collections of Oertijdmuseum De
Groene Poort, Boxtei (thè Netherlands).
Type locality: beach between Nieuwvliet-Bad and
! Cadzand, province of Zeeland (thè Netherlands), dose to
radar and nature reserve ‘de Zwarte Polder’ (Riemslag,
2003, figs. 3, 5).
Description. Carapace subrectangular in outline (Figs.
1-4), length/width (F/W) ratio 1:1.75, widest about half-
way from front. Orbitofrontal margin about two-thirds of
maximum carapace width. Orbits rectangular, about three
times longer than wide. Wide base of thè rostrum covered
with two nodes; rostrum itself not preserved. Short convex
anterolateral margins curving into straight, subparallel
posterolateral margins. Posterior margin straight, slightly
sinuous in centrai part. Regions of carapace very poorly
defined. Deep, almost subcircular epimeral adductor
muscle scars extending into thè cervical furrow anteriorly
and into a short branchiocardiac furrow laterally, forming
two typical hook-shaped depressions. Cervical furrow
directed towards widest part of anterolateral margin. First
thoracic stemite lacking, stemites 2 to 4 are fused; ster-
nites 5, 6 and 7 almost identical in shape and size, sutures
slightly convex posteriorly; seventh stemite barely visible
from ventral side. Abdomen missing; abdominal cavity
extending almost to buccal cavity.
Discussion. So far, thè genus Goniocypoda contained
nine or ten recorded species (Crane, 1981; Crane &
Quayle, 1986; Schweitzer & Feldmann, 2001). In gen¬
erai appearance, G. verheyeni n. sp. seems most closely
related to thè type species, G. edwardsi , from thè middle
Eocene (‘Bartonian’; H. Woodward, 1867, p. 2, pi. 21,
fig. 1) of High Cliff, Hamsphire (England), to G. col¬
limi Crane & Quayle, 1986 (p. 101, pi. 1, fig. 1) from thè
middle Eocene Bracklesham Beds (Lutetian) of Brook,
Hampshire, and to G. elmorensis Crane & Quayle, 1986
(p. 103, pi. 1 , figs. 3, 4) from ?unit 7 of thè Elmore Forma-
tion (Bracklesham Group, middle Eocene) of Hampshire.
The new species differs from thè first-named in having
.
38
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Table 1 - List of post-Paleocene decapod crustaceans currently known from Dutch territory, compiled from
literature data and personal observations.
a L/W ratio of 1:1.75, rather than 1:1. 4- 1.6 (see Tab. 2),
and in showing a more pronounced epimeral muscle scar-
cervical furrow System, plus in showing two nodes at thè
base of thè rostrum.
Goniocypoda collimi differs from G. verheyeni n. sp.
in having a much smaller L/W ratio (1:1.4) as well as in
showing more convex lateral margins and a shallower
and less extensive epimeral muscle scar-cervical furrow
System, while G. elmorensis can be distinguished from
thè new taxon by displaying a ridge on thè orbitofrontal
margin and three tubercles (on thè internai mould) around
thè epibranchial furrow, as well as in having a smaller
L/W ratio (1.6- 1.7, but based on estimated measurements;
see Crane & Quayle, 1986, p. 104).
Goniocypoda verheyeni n. sp. is distinguished from all
other known species of thè genus by its high L/W ratio
(see Tab. 2) and by thè combined features of relatively
large, hook-shaped epimeral muscle scars extending into
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
39
Table 2 - L/W ratios of all species of Goniocypoda known to date. Data from Carter ( 1 898), Remy & Tessier ( 1 954),
Bittner (1893), Glaessner (1933), Glaessner & Rao (1960), Plaziat & Secretan (1971), H. Woodward (1867), Crane
(1981), Crane & Quayle (1986), Schweitzer & Feldmann (2001), Schweitzer et al. (2004) and Guinot (2006).
a straight cervical furrow, of two nodes at thè base of thè
rostrum, of a curved anterolateral margin extending into
a straight posterolateral margin, and of curved stemal
sutures.
A comparison of all species of Goniocypoda on record
to date (Tab. 2) would suggest an increase in carapace
width with stratigraphic time, concomitant with a north-
erly dispersal and several immigration events. Details of
these need to be unravelled; this is beyond thè scope of
thè present paper.
Family Panopeidae Ortmann, 1893
Subfamily Eucratopsinae Stimpson, 1871
Genus Glyphithyreus Reuss, 1859
Type species: Plagiolophus wetherelli Bell, 1858 =
Glyphithyreus affìnis Reuss, 1859, by monotypy, under
ICZN 1999, Artide 67.8.
Glyphithyreus wetherelli (Bell, 1858)
Material: a single phosphatic internai mould (MAB
k.2450) (Fig. 8), collected by Mark Bosselaers (Berchem,
Belgium).
Locality: beach between Nieuwvliet-Bad and Cad-
zand.
Discussion. After thè thorough revision by Karasawa
& Schweitzer (2004), thè genus Glyphithyreus is con-
fined to four species of (?Late Cretaceous)-Paleocene
and Eocene age, placement of Plagiolophus sulcatus
(Beurlen, 1939) in Glyphithyreus being tentative. The
commonest and most widely distributed member of thè
genus is G. wetherelli, with records from thè Paleocene
of Punjab and Baluchistan (Pakistan; Collins & Morris,
1978) and Eocene of northem Germany, southern
England, Belgium, France, Spain and Senegai (Remy
& Tessier, 1954; Plaziat & Secretan, 1971; Moths &
Montag, 2002). Ilyin (2005, pp. 72, 241, pi. 1 1, figs. 4, 5)
recorded G. cf. wetherelli from thè lower Eocene ( Moro -
zovella aragonensis-Araninina pentacamerata Zone) of
Kazakhstan, but these specimens might in fact represent
another species.
Family Carpiliidae Ortmann, 1893
Genus Eocarpilius Blow & Manning, 1996
Type species: Eocarpilius carolinensis, by originai
designation.
Eocarpilius sp.
Material: two specimens in thè C. Riemslag Collec-
tion (Zuidzande, thè Netherlands), previously illustrated
by Fraaye (1998) and Fraaije (2003) as Harpactocarci-
nus sp., plus a single carapace (MAB k.2449) illustrated
here (Fig. 7).
Locality: beach near Nieuwvliet-Bad.
Discussion. Material previously referred to as
Harpactocarcinus sp. by Fraaye (1998) and Fraaije
(2003) is reassigned to thè genus Eocarpilius, which
so far comprised two species from thè middle Eocene
of north Carolina (Blow & Manning, 1996; Feldmann
et al., 1998) and one from thè middle Miocene of Hun-
gary (Miiller, 1984; Feldmann et al., 1998; Schweitzer
et al., 2000; Schweitzer, 2003). The present record
extends thè Eocene range (as documented in eastern
north America) of thè genus to northwest Europe, and
suggests that dispersal during thè Eocene across thè
Atlantic must have been particularly rapid (see Sch¬
weitzer, 2000, 2003).
Family Geryonidae Colosi, 1924
Genus and subgenus Coeloma ( Coeloma ) A. Milne
Edwards, 1865
Type species: Coeloma vigil A. Milne Edwards, 1865,
by originai designation.
Coeloma ( Coeloma ) balticum Schluter, 1879
Material: a single specimen in thè collections of thè
Natuurhistorisch Museum Rotterdam, NMR 9937-01144.
Localities: having been collected from thè so-called
“basaalconglomeraat” (see Posthumus, 1923; De Neve,
1945) at Ootmarsum, thè present specimen (Figs. 5-6) is
considered to have been reworked from Eocene strata.
40
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Figs. 1-8 - 1-4) Goniocypoda verheyeni n. sp., holotype (MAB k.2440), middle-late Eocene, Nieuwvliet-Bad (Zeeland, The
Netherlands), in orbitai, dorsal, ventral and posterior views, respectively. 5-6) Coeloma ( Coeloma ) balticum Schliiter, 1879, either
remanié Eocene or genuine Oligocene element, from thè so-called ‘basaalconglomeraat’ at Ootmarsum (Overijssel, The Netherlands).
Specimen NMR 9937-01144 in thè collections of Natuurhistorisch Museum Rotterdam, in oblique orbitai and dorsal views, respec¬
tively. 7) Eocarpilius sp. (MAB k.2449), middle-late Eocene, Nieuwvliet-Bad (Zeeland, The Netherlands), in dorsal view. 8)
Glyphithyreus wetherelli (Bell, 1 858) (MAB k.2450), middle-late Eocene, Nieuwvliet-Bad (Zeeland, The Netherlands), in ventral view.
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
41
However, it cannot be ruled out entirely that it does
represent a genuine early Oligocene record of thè spe-
cies; precise collection data for this specimen are want-
ing. Previous records include thè middle-upper Eocene
of Cadzand and Nieuwvliet-Bad (Fraaye, 1998; Fraaije,
2003) and thè middle/upper Eocene-lower Oligocene
of northem Germany (Gramann & Mutterlose, 1975;
Fienau, 1984; Freess, 1992; Polkowsky, 2005).
Discussion. Schliiter ( 1 879, pp. 604-608, pi. 1 8, fìg.
3) recorded Coeloma balticum from thè ‘Unter-Oli-
gocàn an der Ostseekiiste’, on thè basis of specimens
‘aus der “blauen Erde” der Bernsteingrube Palmicken
[s/c]’. As Jagt et al. (2006, p. 368) have recently noted,
thè age of thè ‘Bernsteinformation’ at Yantarnyi, Sam-
land Peninsula (Kaliningrad district, Russia), formerly
Palmnicken in east Prussia, is middle Eocene (Lute-
tian), rather than early Oligocene (see Ritzkowski,
1997). This suggests that thè type material of C. bal¬
ticum is of middle Eocene age; thè species apparently
does range upwards into thè (lower) Oligocene as
records from thè Netherlands and northern Germany
show (Polkowsky, 2005).
Discussion
In comparison with faunas recorded from elsewhere in
Europe, thè co-occurrence in thè beach assemblages from
Cadzand and Nieuwvliet-Bad of members of thè (sub)genera
Coeloma {Coeloma), Eocarpilius, Goniocypoda and Glyphi-
thyreus would appear to be indicative of a middle Eocene
age. However, thè new species of Goniocypoda indicates a
tendency for thè carapace in this genus to become wider with
time, and this, combined with a northerly dispersal during
thè middle Paleogene as well as with mass occurrences of
carpiliids and Coeloma recorded from northem Germany
(see Forster & Mundlos, 1982), cannot completely mie out a
late Eocene date for these assemblages either.
References
Bittner A., 1893 - Decapoden des pannonischen Tertiar.
Sitzungsberichte und Denkschrift der Akademie der
Wissenschaften zu Wien, Wien, 102: 44-55.
Blow W. C. & Manning R. B., 1996 - Preliminary
descriptions of 25 new decapod crustaceans from
thè middle Eocene of thè Carolinas, U.S.A. Tulane
Studies in Geology and Paleontology, New Orleans,
29: 1-26.
Carter J., 1 898 - A contribution to thè palaeontology of
thè decapod Cmstacea of England. Quarterly Journal
of thè Geological Society, London, 54: 15-44.
Collins J. S. H. & Morris S. F., 1978 - New lower Terti-
ary crabs from Pakistan. Palaeontology, London, 2 1 :
957-981.
Crane M. D., 1981 - Hexapod crabs of thè genus Gonio¬
cypoda H. Woodward from thè Upper Eocene of
Hampshire. Zoological Journal of thè Linnean Soci¬
ety, London, 72: 1-19.
Crane M. D. & Quayle J., 1986 - Two new hexapod
crabs of thè genus Goniocypoda Woodward (Crusta-
cea, Decapoda) from thè Hampshire Basin. Tertiary
Research, Leiden, 7: 101-105.
De Neve G. A., 1945 - Coeloma balticum Schliiter uit het
basaal-conglomeraat van het Oligoceen in Oost-Ned-
erland. Geologie en Mijnbouw, Utrecht, 7: 8-10.
Feldmann R. M., Bice K. L., Schweitzer Hopkins C.,
Salva E. W. & Pickford K., 1998 - Decapod crus¬
taceans from thè Eocene Castle Hayne Limestone,
North Carolina: paleoceanographic implications. The
Paleontologica! Society, Memoir, Tulsa, 48: 1-28.
Forster R. & Mundlos R., 1982 - Krebse aus dem Alt-
tertiàr von Helmstedt und Handorf (Niedersachsen).
Palaeontographica, Stuttgart, A 179: 148-184.
Fraaye R., 1998 - Paleogene krabben op het strand van
Cadzand en Nieuwvliet-Bad. In: Gids voor strand-
fossielen van Cadzand en Nieuwvliet-Bad. Haaien- en
roggentanden, schelpen, krabben, slangsterren, zoog-
dierresten. Lindemann T. (ed.). Geode, Amsterdam,
30: 68-72.
Fraaije R. H. B., 2003 - Paleogene krabben op het strand
van Cadzand en Nieuwvliet-Bad. In: Gids voor strand-
fossielen van Cadzand en Nieuwvliet-Bad. Lindemann
T. & Fraaije R.H.B. (eds). Oertijdmuseum De Groene
Poort, Boxtei: 68-72.
Freess W. B., 1992 - Cmstaceen aus den mitteloligozànen
Meeressanden bei Leipzig. Aufschluss, Heidelberg,
43: 361-375.
Glaessner M. F., 1933 - New Tertiary crabs in thè collec¬
tion of thè British Museum. TheAnnals and Magazine
of Naturai History, London, (10) 12: 1-28.
Glaessner M. F. & Rao V. R., 1960 - A new species of
crab from thè early Tertiary Fuller’s Earth deposits
of Kapurdi, Rajasthan, western India. Records of thè
Geological Survey of India, Calcutta, 86: 675-682.
Gramann F. & Mutterlose J., 1975 - Krebsfunde aus dem
Alttertiàr am Sarstedt-Lehrter Salzstock (Dekapoda,
Eozàn, Oligozan, Niedersachsen) (Das nordwest-
deutsche Tertiàrbecken, Beitrag Nr. 17). Berichte der
Naturhistorischen Gesellschaft zu Hannover, Hanno¬
ver, 119: 379-401.
Guinot D., 2006 - Rediscovery of thè holotype of Pae-
duma cylindraceum (Bell, 1859) and description of a
new genus of Hexapodidae (Decapoda, Brachyura).
Zoosystema, Paris, 28: 553-571.
Ilyin I. V., 2005 - Melovye i paleogenovye desiatinogie
rakoobraznye (Crustaceomorpha, Decapoda) zapad-
noii chasti Sevemoi Evrazii. Izdatel ’stvo Moskovskogo
Universiteta, Moskva.
Jagt J. W. M., van Bakel B. W. M., Fraaije R. H. B.
& Neumann C., 2006 - In situ fossil hermit crabs
(Paguroidea) from northwest Europe and Russia.
Preliminary data on new records. Revista Mexicana de
Ciencias Geológicas, 23: 364-369.
Karasawa H. & Schweitzer C. E., 2004 - Revision of
thè genus Glyphithyreus Reuss, 1859 (Cmstacea,
Decapoda, Brachyura, Xanthoidea) and recognition
of a new genus. Paleontologica l Research, Tokyo, 8:
143-154.
Lienau H.-W., 1984 - Die marinen Deckschichten (Mitte-
leozàn-Unteroligozàn) der Helmstedter Braunkohlen
(Niedersachsen, BRD). Documenta naturae, Miin-
chen, 22: 1-120.
42
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Lindemann T. & Fraaije R. H .B., 2003 - Gids voor
strandfossielen van Cadzand en Nieuwvliet-Bad. Oer-
tijdmuseum De Groene Poort, Boxtel.
Martin J. W. & Davis G. B., 2001 - An updated classifica-
tion of thè recent Crustacea. Naturai History Museum
Los Angeles County, Scientific Series, Los Angeles,
39: 1-124.
Moths H. & Montag A., 2002 - Tertiare dekapode Krebse
aus Geschieben und dem Anstehenden Norddeutsch-
lands und Dànemarks. Der Geschiebesammler, Wan-
kendorf, 35: 3-30.
Miiller P., 1984 - Decapod Crustacea of thè Badenian.
Geologica Hungarica, Series Palaeontologica, Buda¬
pest, 42: 1-317.
Plaziat J.-C. & Secretan S., 1971 - La faune de crustacés
décapodes des calcaires à Alvéolines yprésiens des
Corbières septentrionales (Aude). Geobios, Lyon, 4:
117-142.
Polkowsky S., 2005 - Decapode Krebse aus dem oberoli-
gozanen Stemberger Gestein von Kobrow (Mecklen-
burg). Tassados, Schwerin, 1: 1-126.
Posthumus O., 1923 - Over de fauna der phosphaatlagen
in Twente (Beneden-Oligoceen). Bijdragen tot de
kennis der palaeontologie van Nederland, IL Verslag
van de Koninklijke Nederlandsche Akademie voor
Wetenschappen, Haarlem, 32: 367-368.
Remy J.-M. & Tessier F., 1954 - Décapodes nouveaux de
la partie ouest de Sénégal. Bulletin de Société géologi-
que de la France, Paris, (6) 4: 185-191.
Riemslag C., 2003 - Het zoeken van fossielen aan het
strand van Cadzand en de Zwarte Polder. In: Gids voor
strandfossielen van Cadzand en Nieuwvliet-Bad. Lin¬
demann T. & Fraaije, R. H. B. (eds). Oertijdmuseum
De Groene Poort, Boxtei: 24-28.
Ritzkowski S., 1997 - K-Ar-Altersbestimmungen der
bemsteinfuhrenden Sedimente des Samlandes (Palào-
gen, Bezirk Kaliningrad). Metallo, Bochum, 66: 19-
23.
Schluter C., 1879 - Neue und weniger gekannte Kreide-
und Tertiàr-Krebse des nòrdlichen Deutschlands.
Zeitschrift der deutschen geologischen Gesellschaft,
Berlin, 31: 586-615.
Schweitzer C. E., 2000 - Tertiary Xanthoidea (Crustacea:
Decapoda: Brachyura) from thè west coast of North
America. Journal of Crustacean Biology, New York,
20:715-742.
Schweitzer C. E., 2003 - Utility of proxy characters for
classifìcation of fossils: an example from thè fossil
Xanthoidea (Crustacea: Decapoda: Brachyura). Jour¬
nal of Paleontology, Tulsa, 77: 1107-1128.
Schweitzer C. E., 2005 - The genus Xanthilites Bell,
1858 and a new xanthoid family (Crustacea: Deca¬
poda: Brachyura: Xanthoidea): new hypotheses on thè
origin of thè Xanthoidea MacLeay, 1838. Journal of
Paleontology, Tulsa, 79: 277-295.
Schweitzer C. E. & Feldmann R. M., 2001 - Differentia-
tion of thè fossil Hexapodidae Miers, 1886 (Decapoda:
Brachyura) from similar forms. Journal of Paleontol¬
ogy, Tulsa, 75: 330-345.
Schweitzer C. E., Feldmann R. M. & Gingerich P. D.,
2004 - New Decapoda (Crustacea) from thè middle
and late Eocene of Pakistan and a revision of Lobono-
tus A. Milne Edwards, 1864. Contributions from thè
Museum of Paleontology, The University of Michigan,
AnnArbor, 31: 89-118.
Schweitzer C. E., Feldmann R. M., Tucker A. B. &
Berglund R. E., 2000 - Eocene decapod crustaceans
from Pulali Point, Washington. Annals of Carnegie
Museum, Pittsburgh, 69: 23-67.
Woodward H., 1 867 - On a new genus of shore-crab, Goni-
ocypoda edwardsi, from thè lower Eocene of Hamp¬
shire. Geologica! Magazine, London, 4(12): 1-3.
René H. B. Fraaije - Oertijdmuseum De Groene Poort, Bosscheweg 80, 5283 WB Boxtei, The Netherlands.
e-mail: info@oertijdmuseum.nl
Barry W. M. Van Bakel - Schepenhoek 235, 5403 GB Uden, The Netherlands.
e-mail: barry.van.bakel@wolmail.nl
John W. M. Jagt - Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, 621 1 KJ Maastricht, The Netherlands.
e-mail: john.jagt@maastricht.nl
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Alessandro Garassino & Antonio De Angeli
Report of Ranilia constricta (A. Milne Edwards, 1880)
(Brachyura, Raninidae) from thè Tyrrhenian (upper Pleistocene)
of Bovetto (Calabria, S Italy)
The studied specimens were collected in Bovetto
quarry, located at N of Reggio Calabria (Calabria - S
Italy), known for its interesting marine fauna since thè
second half of thè Nineteenth Century. Bovetto series, 45
metres thick, preserves a rich malacological fauna of a
warm climate with many specimens of Strombus bubonius
Lamarck, a typical gastropod from thè Tyrrenhian (upper
Pleistocene) of Mediterranean area. Rare fossil mammals,
as Dama dama cfr. D. tiberina and Hippopotamus cfr. H.
amphibius , ha ve been discovered in Bovetto series. Very
rare decapod crustaceans are sometimes discovered in thè
quarry.
These decapod specimens, three-dimensionally pre-
served, show thè following morphological characters:
carapace subovai, strongly convex from side to side,
flattened, almost straight in midiine. Front slightly
raised. Anterolateral margins of carapace subparallel
and posterolateral convergent. Maximum width of cara¬
pace between lateral spines. Preorbitai, supraorbital, and
extraorbital spines strong and prominent. Anterolateral
margin with one spine prominent and elongate. Orbits
forming inverted V directed obliquely ventral from ros-
trum. Dorsal surface of rostrum smooth. Chelipeds stout
having surface omamented with tubercles and cibate
lines. Movable finger of chela unarmed. Palm of chela
higher than long with ventral margin terminating in a
strong spine, and with opposable margin not armed with
teeth. Merus of cheliped with blunt dorsal spine.
The above-mentioned morphological characters are
typical of Ranilia constricta (A. Milne Edwards, 1880) to
which thè studied specimens are assigned. At present, this
species is widespread in thè western Atlantic (to Florida
Straits and Yucatan Channel, Cuba, off Barbados, and
Brazil), Central Atlantic (Ascension Island), and eastem
Atlantic (from Senegai to Congo). At present, only one
species is recognised in thè fossil record, R. punctu/ata
Beschin, Busulini, De Angeli & Tessier, 1988, from thè
lower Eocene (Ypresian) of Valle del Chiampo (Vicenza,
N Italy) (Beschin et al. 1988), since Feldmann & Max¬
well (1990) assigned R. pororariensis Glaessner, 1980,
from thè upper Eocene of New Zealand to Laeviranina
Lòrenthey & Beurlen, 1929. Miiller (1993) ascribed some
specimens from thè Pliocene of Spain to Ranilia , but this
ascription is stili uncertain. Porteli & Roger (2003) pub-
lished a peer-reviewed abstract on thè frog crab Ranilia
from thè Pliocene of Florida (United States), and, as
reported by Porteli & Agnew (2004), a new fossil species
will be described.
Notopella Lòrenthey & Beurlen, 1929 ( N . vareolata
Lòrenthey & Beurlen, 1929, type species) was
synonymised to Ranilia by Glaessner (1969) and Miiller
& Collins (1991), while Via Boada (1965, 1969), Jagt
et al. (1993), and Tucker (1998) considered Notopella
Lòrenthey, 1929, a valid genus within thè subfamily
Notopodinae.
References
Beschin C., Busulini A., De Angeli A. & Tessier G.,
1988 - Raninidae del Terziario berico-lessineo (Italia
settentrionale). Lavori - Società Veneziana di Scienze
Naturali, Venezia, 13: 155-215.
Feldmann R. M. & Maxwell P. A., 1990 - Late Eocene
Decapod Crustacea from North Westland, South
Island, New Zealand. Journal of Pa/eonto/ogy, Law¬
rence, 64 (5): 779-797.
Glaessner M. F., 1969 - Crustacea Decapoda. In: Treatise
on Invertebrate Paleontology. Arthropoda 4 (2), Geo¬
logica I Society of America and University of Kansas,
Lawrence: R399-R533, R626-R628.
Jagt J. W. M„ Collins J. S. H. & Fraaije R. H. B., 1993 -
A new Early Palaeocene genus of raninid crab
(Crustacea, Decapoda) from Denmark, Southern
Sweden and The Netherlands. Contributions to
Tertiary and Quaternary Geology, Leiden, 30 (3-4):
177-182.
Miiller P., 1993 - Neogene Decapod Crustaceans from
Catalonia. Scripta Musei Geologici Seminarii Barci-
nonensìs, Barcelona, 225: 1-39.
Miiller P. & Collins J. S. H., 1991 - Late Eocene coral-
associated decapods (Crustacea) from Hungary.
Contributions to Tertiary and Quaternary Geology,
Leiden, 28 (1-2): 47-92.
Porteli L. & Agnew J. G., 2004 - Florida Fossil lnverte-
brates. Pliocene and Pleistocene decapod crustaceans.
Florida Paleontologica/ Society, 4: 1-10.
Porteli L. & Roger W., 2003 - Exceptional preservation
and concentration of whole-body Ranilia (Decapoda:
Raninidae) in thè Pliocene Intracoastal Formation of
Florida. Geologica! Society of America, Southeastern
Section, Abstracts with Programs, 35 (1): 68-69.
Tucker A. B., 1998 - Systematics of thè Raninidae (Crus¬
tacea: Decapoda: Brachyura) with accounts of three
new genera and two new species. Proceedings of thè
44
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Biologica I Society of Washington, Washington, 111
(2): 320-371.
Via Boada L., 1965 - Raninidos fósiles de Espana. Con-
tribución al estudio paleontològico de la familia “Rani-
nidae” (Crustàceos decàpodos). Boletino Instituto Ge¬
ologico y Mineralogico de Espana, Madrid, 76: 3-43.
Via Boada L. 1969 - Crustàceos decàpodos del Eoceno
espandi. Pirineos, Barcelona, 91-94: 1-479.
Fig. 1 - Ranilia constricta (A. Milne Edwards, 1880). A) Lateral view. B) Frontal view. C) Dorsal view. D) Reconstruction.
Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: agarassino@libero.it
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
3"1 Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Alessandro Garassino, Antonio De Angeli & Giovanni Pasini
New decapod assemblage from thè Upper Cretaceous
(Cenomanian-Turonian) of Gara Sbaa, southeastern Morocco
The previous reports of decapods from thè Upper
Cretaceous of Morocco are limited to two discoveries of
“pinces de crabes” respectively from thè Cenomanian of
thè Akrabou-section of Ziz Formation (CI unit), and from
thè upper Cenomanian of thè section of Goulmina (C2
unit - upper part), both located in thè Errachidia region
(Ettachfini & Andreu, 2004). Dutheil (1999) reported thè
presence of decapod crustaceans in thè faunal assemblage
of thè locality “dose to Gara Sbaa”, in thè upper part of
thè Kem Kem beds (Cenomanian in age). Garassino et
al. (2006) described Cretapenaeus berberus, a freshwa-
ter decapod crustacean, on thè specimens gathered by
Dutheil from this locality, located more to thè north of
Gara Sbaa.
The recent discovery of anomurans, brachyurans, and
macrurans from a new locality, excavated by field research
carried out by one of thè authors (G. P.) for thè Sezione di
Paleontologia degli Invertebrati del Museo di Storia Natu¬
rale di Milano, permitted an increasing of thè carcinologie
knowledge about these systematic groups. The studied
specimens were discovered in new deposits at thè top of
Gara Sbaa escarpment, located in SE Morocco, along thè
Hamada of Kem Kem, dose to thè Algerian border. These
fossiliferous levels are known in literature as Kem Kem
beds (Sereno et al., 1996) from thè Upper Cretaceous. The
sediments are exposed from NE to SE for an extent of 250
km and they are marked to thè N by Tafilalt area, to thè
E by Hamada of Guir, to thè S by Hamada of Kem Kem,
and to thè W by Precambrian formations and Paleozoics
of Anti-Atlas. The studied area, located SW of Taouz, 26
km S/SW of Taffaout, along thè Oued Sbaa, is formed by a
small sedimentary series, located at thè top of a small mesa,
known as Gara Sbaa. The new sedimentary levels, cover-
ing thè upper unit of Kem Kem beds, lie directly on thè
Cenomanian-Turonian limestones. These levels, having a
small extent (about 500 m2) and 1.80 m thick, show at thè
bottoni sublithographic laminated limestones (60/70 cm
thick), including thè faunal assemblage, thè subject of this
study. In addition to decapod crustaceans, many inverte¬
brate and vertebrate taxa were discovered in these levels:
xiphosurans (first report in N Africa), isopods, peracarids,
rare insects (Orthoptera and Hemiptera), traces of marine
worms, rare pelagic crinoids (Comatulidae), rare teeth of
chondrichthyes, and many well-preserved specimens of
actinopterygians. Finally, well-preserved leafy branches
and leaves were also discovered. All fossils have a high
level of preservation resembling thè taphonomy of other
“ lagerstàtten ” quarries.
Some specimens of macrurans are assigned to Cre¬
tapenaeus berberus Garassino, Pasini & Dutheil, 2006,
recently described from another locality and unit of thè
Kem Kem beds.
Many specimens of glypheids are assigned to a new
species of Glyphea v. Meyer, 1 835 (Garassino et al., work
in progress). At present, w e recognize live species from
thè Upper Cretaceous: G. bohemica Fritsch & Kafka,
1887 (Turonian - Boemia), G. cretacea McCoy, 1854,
and G. willetti (Woodward, 1878) (Cenomanian - Great
Britain), G. damesi Garassino, 2001 (Cenomanian - Leba-
non), and G. foresti Feldmann & de Saint Laurent, 2002
(Cenomanian - Australia) which differ from thè new spe¬
cies by thè presence of two or three carinae in gastric and
antennal regions (only one in thè new species) and for thè
smooth dorsal and ventral margins of rostrum (tuberculate
margins in thè new species).
Some specimens of galatheids are assigned to a new spe¬
cies of Galathea Fabricius, 1793, Paragalathea Patrulius,
1960, and to a new genus within thè family Galatheidae
Samouelle, 1819 (Garassino et al., work in progress). Even
though thè studied specimens belonging to a new species
of Galathea partially preserve thè rostrum, they show
morphological characters comparable with this genus, such
as thè transverse cristae of thè carapace and thè triangular
shape of thè rostrum. The new species differ from thè other
fossil species by thè carapace being weakly developed in
length, with almost rectilinear cristae continuous to thè lat-
eral margins, and exhibiting very developed spines of thè
rostral margins. The studied specimens belonging to a new
species of Paragalathea differ from thè other fossil spe¬
cies by having thè carapace wider than long and thè dorsal
granules weakly evident and raised. The studied specimens
belonging to a new genus have morphological characters
that do not resemble those of thè other galatheids known
to date. In fact, this new genus has thè anterior part of
thè carapace strongly restricted, thè fronto-orbital margin
wide, about 0.55 of thè maximum width of thè carapace;
thè antero-lateral margins concave and strongly divergent
to thè small spine of thè epibranchial angle; thè lateral mar¬
gins of thè carapace with small teeth; and trans verse cristae
of thè median and posterior parts of thè carapace that do not
reach thè lateral margins.
Some specimens of brachyurans are ascribed to
Corazzatocarcinus hadjoulae (Roger, 1946) and Telamo-
nocarcinus gambalatus Larghi, 2004, recently described
by Larghi (2004) from thè Upper Cretaceous (Cenoma¬
nian) of Lebanon.
Finally, some specimens of brachyurans are ascribed
to a new genus and new species (Garassino et al., work in
progress). These specimens have thè carapace as long as
wide with dorsal regions weakly marked, front with four
lobes, small orbits, supraorbital margin strongly concave,
and anterolateral margin with four spines. The sternites
are really typical, having an ovai elongate shape and
narrow anterior part.The sutures 1/2 and 3/4 are complete,
46
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Fig. 1 - A) Glypheidae n. sp., MSNM i268 1 8. B) Galatheidae n. sp., MSNM Ì26827. C) Galatheidae n. sp., MSNM Ì26862. D) Gala-
theidae n. gen., n. sp., MSNM Ì26856. E-F) Bellidae? n. gen., n. sp., MSNM Ì26858 (dorsal view), MSNM Ì26831 (ventral view).
3“ SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
47
thè sutures 4/5, 5/6, and 7/8 are interrupted in thè median
part, thè suture 6/7 is complete, and thè sutures 5/6, 6/7
and 7/8 are strongly curved posteriorly; stemo-abdominal
cavity weak, deep; vulvae located after suture 5/6. Ambu-
latory legs II-V elongate with tapering dactyli.
The new genus does not show strict correlations
with any crabs known to date. Therefore, we justify thè
description of a new genus belonging to an indeterminate
family within thè Eubrachyura.
References
Dutheil D. B., 1999 - An overview of freshwater fìsh
fauna from thè Kem Kem beds (Late Cretaceous:
Cenomanian) of southeastem Morocco. In: Mesozoic
Fishes 2 - Systematics and thè fossil Record. Arratia
G. & Schultze H.-R (eds). VerlagDr. Friederich Pfeil :
553-563.
Ettachfini El. M. & Andreu B., 2004 - Le Cénomanien et
le Turonien de la Piate-forme Préafricaine du Maroc.
Cretaceous Research , 25: 277- 302.
Garassino A., Pasini G. & Dutheil D. B., 2006 - Cretape-
naeus berberus n. gen., n. sp. (Crustacea: Decapoda:
Penaeidae) from thè Late Cretaceous (Cenomanian)
of southeastem Morocco. Atti della Società italiana di
Scienze naturali e del Museo civico di Storia naturale
in Milano, Milano, 147 (1): 3-17.
Larghi C., 2004 - Brachyuran decapod crustacea from thè
Upper Cretaceous of Lebanon. Journal of Paleonto-
logy, Lawrence, 78 (3): 528-541.
Sereno P. C., Dutheil D. B., Iarochène M., Larsson H. C.
E., Lyon G. H., Magwene P. M., Sidor C. A., Varric-
chio D. J. & Wilson J. A., 1996 - Predatory dinosaurs
from thè Sahara and Late Cretaceous faunal differen-
tiation. Science, 272: 986-991.
Alessandro Garassino - Museo Civico di Storia Naturale, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: agarassino@libero.it
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 1 5, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
Giovanni Pasini - Museo Civico dei Fossili di Besano, Via Prestini 5, 21050 Besano (Varese), Italy.
e-mail: juanaldopasini@tiscali.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Alice Giannetti, Paolo Monaco, Jesus E. Caracuel, Jesus M. Soria &
Alfonso Yébenes
Functional morphology and ethology of decapod crustaceans
gathered by Thalassinoides branched burrows in
Mesozoic shallow water environments
Every part of a burrow has an important and specific
function with regard to thè ecology and trophism of thè
burrower. It goes from thè guarantee of thè oxygena-
tion and irrigation of thè tunnels to thè protection of thè
organism or thè storing of organic material for feeding
(Nickell & Atkinson, 1995). Regarding modem thalassi-
noidean burrows, presence of surface mounds, vertical
development of tunnels, geometry of thè section of tun¬
nels and horizontal galleries, presence of chambers and
of organic debris within thè burrows, number and type
of apertures on thè seafloor, presence of exhalant tunnels
and Y- or U-shaped burrow in vertical section, are gen-
erally considered as thè most important characteristics
for ethological characterization of thè burrow and of
thè trace-maker (Nickell & Atkinson, 1995). Some of
these features can be recovered also in thè trace fossil
Thalassinoides, usually considered a fossil counterpart
of modem thalassinoidean burrows, and they allow
one to formulate hypotheses about thè trophism of thè
trace maker. This can be particularly useful in those
cases in which body-fossil remains of thè burrowers
are completely absent and a direct comparison between
thè morphology of ancient and modern organisms is not
possible. Ecologie considerations are here applied to
Pliensbachian and Albian Thalassinoides developed in
thè lagoonal deposits of thè Trento carbonate platform
(Rotzo Member, Calcari Grigi Formation, northern
Italy) and in thè outer shelf deposits of thè Sàcaras For¬
mation (Serra Gelada, Alicante Province southeastem
Spain), respectively. Different morphotypes of Tha¬
lassinoides were identifìed in these two areas, mainly on
thè basis of dimensions of thè branches, diameter of thè
turning chamber and development and geometry of thè
maze. Even if Thalassinoides like burrows are nowadays
produced also by worms and fishes, we consider crusta¬
ceans to be thè most probable trace makers of thè studied
Thalassinoides. For a detailed discussion of thè geologi¬
ca! setting, palaeonvironmental models, and ichnology
conceming thè studied areas see Avanzini (1998), Gian¬
netti (2004), Giannetti & Monaco (2002), Masetti et al.
(1998), Masetti & Posenato (2002), Monaco (2000a,
2000b), Monaco & Garassino (2001), Monaco & Gian¬
netti (2001, 2002), Winterer & Bosellini (1981), and
Zempolich (1993) for thè Calcari Grigi Formation, and
Caracuel et al. (2002), Castro (1998), Giannetti et al.
(2005), Monaco et al. (2005), Vilas et al. (1982), and
Yébenes (1996) for thè Sàcaras Formation and refer-
ences therein.
In thè Rotzo Member, three types of Thalassinoides
suevicus Rieth 1932 ( Thalassinoides type I to type III,
from thè smallest to thè largest form) and another form
type of Thalassinoides isp. (type IV) were identifìed.
Thalassinoides suevicus type I is a small burrow, with
a diameter ranging from 2 cm to 5 cm. Branches are thin
and circular in section. Turning chambers are not par¬
ticularly developed. Mazes are regular and geometrical
in shape and developed only on thè horizontal piane.
Burrows are mud-fìlled and their surface is smooth. When
these burrows are in tiered with thè other forms of Tha¬
lassinoides, they occupy thè upper tier.
In Thalassinoides suevicus type II, thè diameter of
branches ranges from 5 cm to 1 0 cm and diameter of tum-
ing chambers ranges from 6 cm to 10 cm. Branches are
long and form geometrical, sometimes pentagonal mazes.
The extemal surface is smooth and lining is absent. Mud-
stone or bioclastic wackestone fills thè burrows. Bioclasts
can be minutely fragmented and grouped in thè internai
part of thè burrow. This is thè most frequently observed
type of Thalassinoides in thè studied sections.
Thalassinoides suevicus type III is one of thè largest
Thalassinoides form. Its diameter ranges between 10 cm
and 1 6 cm, and thè diameter of thè turning chamber is up
to 22 cm. Branches are squat with respect to thè morphol¬
ogy of thè whole burrow and bifurcate at regular angles.
The outer surface is smooth and thè filling is made of
bioclastic wackestones/packstones. Mottled concentra-
tions of minutely fragmented bioclasts can be present in
thè centrai part of thè burrow, in correspondence to thè
lumen. This form always occupies thè deepest tiers.
In Thalassinoides type IV thè diameter of thè branches
can reach 16 cm. The outer surface is mammillary, with
small mounds distributed without particular orientation
with respect to thè burrow. Branches are short, subcircu-
lar in section and mazes are irregularly developed. On thè
outer surface, crinoids and other bioclasts are frequently
grouped. Bioclastic wackestone fills thè burrows.
Other cylindrical burrows, indicated as Thalassi-
noidesl isp., exhibit simple tubes up to 80 cm in length,
are gently arched, 6 to 8 cm in diameter, and show short
aborted branches, without turning chambers. These
traces are very similar in horizontal piane view to those
illustrated by Bromley (1990). These atypical, branched
tunnels are very different from I-II-III-IV types described
above, and are analogous to some burrows of stomatopods
from thè Seychelles Islands. For their morphological and
taphonomic characteristics these simple cylindrical tubes
3™ SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
49
are not closely related to burrows of shrimps or other
decapods, but resemble to those of camivorous crusta-
ceans (stomatopods, Malacostraca), indicated as active
predators, now populating thè shallow waters mainly of
tropical and subtropical seas.
In thè Sàcaras Formation, we identified three types
of Thalassinoides suevicus Rieth 1932 ( Thalassinoides
type A to type C) and another poorly branched form
( Thalassinoides type D), resembles thè Thalassinoides ?
isp. described for thè Rotzo Member, while thè type IV is
very rare.
Thalassinoides suevicus type A is a small burrow,
with a maximum diameter of 5 cm. Tuming chambers are
present but not particularly developed. It does not show
laterally developed mazes, but only burrows with short
branches developed only on thè horizontal piane. Bur¬
rows are mud-filled and their surface is smooth. Similarly
to thè Rotzo examples, these burrows can be frequently
observed tiered with thè other forms of Thalassinoides
and they are placed in thè most superficial layers.
In Thalassinoides suevicus type B, branch diameter
ranges from 6 cm to 8 cm, tuming chambers, can reach a
diameter of 9 cm. Regular mazes can be well developed
on thè horizontal piane. The outer surface is smooth,
lining is absent and fìlling is homogenous and made up of
bioclastic wackestone.
Thalassinoides suevicus type C is thè largest Thalassi¬
noides form, with a mean diameter of around 8 cm, and
up to 12 cm in size for thè tuming chamber. When tier-
ing is present, this form is always located in thè deepest
layers. Filling is homogeneous and made up of bioclastic
wackestone.
Thalassinoides isp. type D is a very large, horizontally
developed burrow, following an irregular zigzag path. It
is up to 1 m long, with a diameter ranging between 10
cm and 14 cm and a circular to slightly elliptical section.
It does not show true branching, but only short protru-
sions, extending in correspondence of thè bending point
of thè burrow. This trace resembles Thalassinoides ? isp.
of Calcari Grigi. Filling is generally homogeneous and
is made up of fìne-grained calcarenites or by bioclastic
wackestone/packstone, according to thè lithology of thè
host rock.
The following interpretation of thè trophism of thè
trace makers and of thè ecological significance of thè
different Thalassinoides forms are based on thè model
proposed by Nickell & Atkinson (1995; fig. 3, p. 195)
for modem Thalassinoides-Mkt burrows, considering thè
main features common for both, modem and fossil bur¬
rows. The 12 diagnostic features suggested by Nickell and
Atkinson (1995) and their ecological significance are here
briefly examined.
1) Surface mounds. They are produced by thè trace-
maker removing sediment from thè burrow and transport-
ing it to thè surface. Surface mounds formed after thè
construction of thè burrow are indicative of deposit feed-
ing trophic mode. Cone-shaped, muddy mounds, locally
crossed by narrow vertical shafts are represented by a
few specimens in thè Rotzo Member and are completely
absent in thè Sàcaras Formation, maybe due to thè higher
intensity of bottom currents.
2) Horizontal mazes (it replaces thè “tightly layered
lattice” feature of Nickell & Atkinson, 1995). They sug-
gest thè intense exploitation of thè sediment by a deposit
feeder. Although all thè studied Thalassinoides are hori¬
zontally developed, this feature is particularly character-
istic of Thalassinoides type II, forming geometrical mazes
widely developed at thè base of thè beds. As suggested by
Suchanek et al. (1986), laterally extended mazes would
allow thè trace makers to maximize thè capture of organic
material when thè substratum has a low nutritional value.
This probably could be related to different rates of nutri-
ent input within thè sediment in thè various phases of
evolution of thè lagoon.
3) Deep burrows. Depth of thè burrow development
within thè substratum can indicate thè importance of sur¬
face sediments as a nutritional source for thè trace-maker.
The deeper thè maze is developed within thè sediment, thè
less thè trace maker depends directly on organic materials
and current destruction present on thè seafloor. Due to thè
absence of vertical tunnels and to thè vertical distribution
of thè different Thalassinoides types within thè beds, we
have hypothesized that thè studied Thalassinoides, and in
particular thè smallest forms, were probably developed
at a shallow depth within thè substratum. This would
indicate a quite strong dependence of thè trace makers on
organic material present on thè seafloor (surface deposit
feeders or omnivorous scavengers). We must consider thè
possibility that parts of burrows were eroded by bottom
currents and depth of burrowing within thè substratum
could have been therefore underestimated. This consider-
ation is particularly important in thè Serra Gelada section,
where burrows are developed in a relative high-energy
environment.
4) Sub-circular tunnel cross section. According to
Nickell & Atkinson (1995), sub-circular tunnels occur in
established burrows, after thè phase of exploitation, and
are thè result of crustacean movement. Ovai or irregu¬
lar shapes are less suitable for efficient water flow than
a perfectly circular cross-section (Nickell & Atkinson,
1995). The narrowing in cross section, as occurs in some
Thalassinoides ? isp. (such as in some modem burrows of
stomatopods), is another characteristic which is not ideal
to water flow. According to this hypothesis, thè occupant
of thè burrow would be strongly dependent on thè sur¬
face sediments as a nutritional source and less to current
flows. With thè exception of Thalassinoides type IV, all
thè studied Thalassinoides both in thè Calcari Grigi and
in thè Sàcaras Formation have a sub-circular cross section
or narrowing, thus confìrming thè hypothesis suggested
by thè other characters that thè primary nutritional source
was thè organic material present on thè seafloor rather
than that transported passively by current flow within thè
tunnels.
5) Chambered burrows. Chambers can have different
purposes: they can be used for sub-surface deposit feed-
ing or to store coarse grained material and fragmented
bioclasts, found within thè substratum or coming from thè
sea-floor. Another purpose maybe thè change thè direc¬
tion of crustacean by 180°, by performing a somersault
followed by a roll around thè body axis, as observed in
modem examples of thalassinoideans. Therefore, thè sub-
spherical shape of chambers maybe important to recognize
behaviour pattems, in thè relationship with charateristics
of thè substrate (Monaco & Giannetti, 2002). In Thalassi¬
noides type III, type IV and secondarily in Thalassinoides
type II of thè Calcari Grigi Formation and type B and C of
thè Sàcaras Formation, tuming chambers are particularly
50
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
developed and large. Stored coarse grained material such
as fragments of crinoids, coated grains and large bivalves,
algae and foraminifers were sometimes observed, in par-
ticular in Thalassinoides type IV.
6) Organic detritus in burrows. Organic detritus in bur-
rows can be thè result of scavenging on thè surface, but
also of passive fading of material within an open burrow.
In thè fossil record, thè different origin of organic detri¬
tus can be sometimes identified through taphonomic and
sedimentologie criteria. In modem Thalassinoides- like
burrows, organic material is also used by thè crustacean
to cultivate micro-organisms for nutritional purposes.
Organic detritus in burrows is particularly abundant in
Thalassinoides type III and type IV.
7) Oblique tunnels were very rarely recorded in thè
studied sections (in some Thalassinoides ? isp.) and there-
fore they are not considered here.
8) Burrow openings, tubular tempestites. In thè studied
sections, burrow openings can be directly observed only
rarely. Their existence can be deduced by thè presence
of tubular tempestites inside thè bed, but their number,
dimensions and density cannot be defined. The presence
of many burrow openings would indicate a continuous
and necessary contact of thè trace maker with thè seafloor,
for deposit feeding or scavenging.
9) Funnel-shaped openings. According to Nickell &
Atkinson (1995), thè presence of funnel-shaped openings
would facilitate thè capture of prey or any other nutri¬
tional particles, as they fall down to thè sides of thè funnel
and into thè burrow. In thè studied section, this feature is
represented only by a specimen from thè Rotzo Member.
The funnel is asymmetric in shape, thè back being nearly
vertical and thè front being flattened probably due to thè
continuous movement of thè trace maker. The bottoni of
thè funnel is connected to a short vertical tunnel, which is
not directly related to a preserved Thalassinoides maze.
10) Narrow exhalant shaft. The presence of a narrow
exhalant shaft indicates thè necessity of current genera¬
tion. This feature was observed only in one cone-shaped
mound in thè Rotzo Member, but its origin is uncertain.
11) U- or Y- shaped burrows in vertical section. This
feature, typical of many modem thalassinoidean burrow Sys¬
tems (Bromley, 1 990), was not observed in thè studied fossil
Thalassinoides and therefore it is not considered here.
12) Circular tunnel cross section. Their presence
would facilitate thè water flow within and through thè
burrow. Due to thè importance of a Constant water flow
through thè burrow, circular tunnel cross sections are
present also in burrows produced by primarily deposit
feeders and therefore this cannot be considered diag-
nostic of a particular trophic mode (Nickell & Atkinson,
1995). In thè studied sections this feature is only rarely
represented in Thalassinoides type II and in thè largest
Thalassinoides type III, mainly where mazes are well
preserved and regularly developed.
In summary, features representing sediment processing
and Storage of material for nutritional and maintenance
purposes are thè most common in thè Thalassinoides of
thè Rotzo Member.
The only characteristic shared by thè five Thalassi¬
noides forms is thè horizontal development of thè bur¬
rows. Branches with a sub-circular section are typical
of Thalassinoides suevicus type I, type II, and type III.
Enlarged chambers are absent in Thalassinoides suevicus
type I and Thalassinoides ? isp., present in type II and type
III and well developed in type IV.
According to thè scheme proposed by Nickell &
Atkinson (1995), thè most commonly observed features
in all thè Thalassinoides of thè Calcari Grigi Formation
indicate intense sediment processing. In Thalassinoides
suevicus type II, type III, and especially in thè type IV,
Storage of organic material (chambered burrows and
organic detritus in burrows, characters 5 and 6) is also
evidenced. The presence of organic detritus is represented
by coarse-grained to fine-grained cmshed bioclasts within
Thalassinoides suevicus type II, type III, and type IV. It
indicates both biogenic sorting and Storage of bioclas-
tic debris coming from thè seafloor and fallen down or
trasported into thè burrow. The locai concentration of
coarse-grained fragmented shells in Thalassinoides type
IV, is probably due to thè need of thè trace maker to main-
tain other parts of burrow free from debris.
Vertical tunnels and apertures on thè seafloor (char¬
acters 8, 9 and 12) are very rarely observed and it is
sometimes impossible to state if vertical tunnels are not
preserved due to sedimentologie or diagenetic processes
or if they were really poorly developed in thè burrows.
Actually, we have no evidences of deep burrows with a
U- or Y- shaped vertical morphology. We have hypoth-
esized, therefore, that mazes were burrowed on horizontal
planes only at shallow depth in thè substratum. The few
evidences of vertical tunnels and of exhalant shafts are
difficult to relate to Thalassinoides mazes. The presence
of tubular tempestites inside thè beds proves thè existence
of direct apertures on thè seafloor, even if only thè hori¬
zontal part of thè filled burrows can be usually observed.
The only clearly observable vertical element has a circu¬
lar section (character 12), but it is an isolated feature. This
tunnel closely resembles thè burrows of modem crusta-
ceans and therefore we have considered it separately. In
thè category of thè vertical elements, we have considered
also inhalant and exhalant shafts, represented in thè Rotzo
Member only by an unclear example. Actually, shafts
are so small that they could be produced also by worms.
These rare vertical elements indicate thè presence of sus-
pension feeders, whose primary nutritional source is thè
water column, and of surface deposit. feeders. According
to thè scheme proposed by Nickell & Atkinson (1995), all
thè studied Thalassinoides belong to thè category of thè
sub-surface deposit feeders (searching for organic parti¬
cles within thè substratum) and secondarily to that of thè
omnivorous scavengers (Vaugelas, 1990), which ingest
organic debris present on thè seafloor and deriving from
other animals and algae.
Similar interpretations can be proposed for thè Tha¬
lassinoides studied in thè Sàcaras Formation, even if
only four of thè twelve characters described by Nickell
& Atkinson (1995), namely those related thè horizontal
development of thè burrow (characters 2-5), are well
preserved. The problem concerns thè vertical develop¬
ment of thè burrows, because we have neither direct
nor indirect proofs of thè existence of vertical tunnels
(except Ophiomorpha specimens of calcarenites at thè
top of parasequences of Serra Gelada, see Monaco et
al., 2007) and of thè type and number of apertures on
thè seafloor. Anyway, we can hypothesize thè pres¬
ence at least of some vertical tunnels linking thè big-
gest Thalassinoides, placed in thè deepest tiers within
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
51
thè sediment, to thè seafloor. Intense bioturbation of
thè more superfìcial tiers probably obliterated them.
All thè studied Thalassinoides are horizontally devel-
oped ( horizontal mazes, character 2) and show a sub-cir-
cular cross section (character 4), both characters indica¬
tive of intense sediment processing.
Presence of enlarged chambers (as in Thalassinoides
suevicus type B and in Th. suevicus type C, character
5) could indicate Storage of organic material for feed-
ing purposes or changes in thè direction of locomotion.
Concentration of skeletal remains (character 6) is rarely
observed, due to thè high bioclastic content of thè host
rock and of thè burrow filling. Because of thè absence of
vertical tunnels, we cannot formulate hypotheses about
conditions of irrigation and importance of thè water
column for nutritional purposes (characters 7-12). It is
evident that burrowers were closely dependent on nutri-
ents present within and on thè substratum as thè main
feeding source. Therefore, they can be included in thè
trophic categories of thè sub-surface deposit feeders and
secondarily in those of thè surface deposit feeders and of
thè omnivorous scavengers.
This is only a first step in thè ecological analysis of thè
Thalassinoides trace fossils and of thè trophism of their
trace-makers. However, an approach which considers thè
functional morphology of thè parts making up modem
Thalassinoides- like burrows can be useful for understand-
ing of thè ecological meaning of ancient burrows produced
by extinct crustaceans and can give further information
for a more complete paleoenvironmental reconstruction
(Monaco et al., 2007). However, this method has to be
refined, elaborating a detailed model which compares dif-
ferent types of Thalassinoides in different environmental
contexts and considering also thè functional morphology
of modem branched burrows produced not only by crus-
tacean decapods but also by other organisms.
References
Avanzini M., 1998 - Resti di rettili continentali dal
Giurassico inferiore della piattaforma di Trento (Italia
settentrionale). Studi Trentini di Scienze Naturali -
Acta Geologica, Trento, 73: 75-80.
Bromley R. G. C., 1990 - Trace fossils, biology and
taphonomy. Special topics in paleontology. Unwin
Hyman Ltd, London: 1-280.
Caracuel J. E., Monaco P., Yébenes, A. & Giannetti A.,
2002 -Trazas afines a Imhrichnus wattonensis Hallam
de edad Albiense en el Prebético de Alicante (Serra
Gelada) Geogaceta, Zaragoza, 31: 171-174.
Castro J. M., 1998 - Las Plataformas del Valanginiense
superior- Albiense superior en el Prebético de Alicante.
Unpublished Ph.D. Thesis, University of Granada.
Giannetti A., 2004 - Analisi tafonomica ed ichnologica
delle facies a Thalassinoides del Mesozoico Tetideo.
Unpublished Ph. D. Thesis. University of Pemgia.
Giannetti A., Caracuel J. E., Monaco P., Soria J. M. &
Yébenes A., 2005 - Sedimentologia, tafofacies e icno-
coenosis de las parasecuencias albienses de rampa
carbonatada extema en el Prebético de Alicante (Serra
Gelada). Geotemas, Salamanca, 8: 57-62.
Giannetti A. & Monaco P., 2002 - Burrow-Decreasing
Upward Parasequence (BDUP): A case study from thè
Lower Jurassic Trento Carbonate Platform (Southern
Alps). Rivista Italiana di Stratigrafia e Paleontologia,
Milano, 110(1): 77-85.
Masetti D., Claps M., Giacometti A., Lodi, P. & Pignatti
P., 1998 - I Calcari Grigi della Piattaforma di Trento
(Lias inferiore e medio, Prealpi Venete). Atti Ticinensi
di Scienze della Terra, Pavia, 40: 139-183.
Masetti D. & Posenato R., 2002 - The Trento Plateau and
thè Belluno Basin. Field trip B5. Stop.3-Valbona. The
Middle Liassic Lithiotis facies of thè Trento Platform
(Calcari Grigi, Membro di Rotzo). General Field
Trip Guidebook, 6th International Symposium on thè
Jurassic System, Palermo (Italy): 283-288.
Monaco P., 2000a - Biological and physical agents of
shell concentrations of Lithiotis facies enhanced by
microstratigraphy and taphonomy, Early Jurassic,
Gray limestones Formation, Trento area (Northern
Italy). GeoResearch Forum, Zurich, 6: 473-485.
Monaco P., 2000b - Decapod burrows ( Thalassinoides ,
Ophiomorpha ) and crustacean remains in thè Calcari
Grigi, lower Jurassic, Trento platform (Italy). In: lst
Workshop on Mesozoic and Tertiary decapod crus¬
taceans. Studi e Ricerche - Associazione Amici del
Museo - Museo Civico “G. Zannato”, Montecchio
Maggiore: 55-57.
Monaco P., Caracuel J. E., Giannetti A., Soria J. &
Yébenes A., 2007 - Thalassinoides and Ophiomorpha
as cross-facies trace fossils of cmstaceans from shal-
low-to-deep-water environments: Mesozoic and Terti¬
ary examples from Italy and Spain. In: 3rd Symposium
on Mesozoic and Cenozoic Decapod Cmstaceans.
Museo di Storia Naturale di Milano, May 23-25, 2007.
A. Garassino, R. M. Feldmann & G. Temzzi (eds.).
Memorie della Società italiana di Scienze naturali e
del Museo civico di Storia naturale di Milano, Milano,
XXXV (II): 79-82.
Monaco P. & Garassino A., 2001 - Burrowing and body
fossil of decapod cmstaceans in thè Calcari Grigi,
Lower Jurassic, Trento platform (Italy) Geobios,
Lyon, 34 (3): 291-301.
Monaco P. & Giannetti A., 2001 - Stratigrafia tafonomica
nel Giurassico Inferiore dei Calcari Grigi della Piatta¬
forma di Trento. Pavia. Atti Ticinensi di Scienze della
Terra, Pavia, 42: 175-209.
Monaco P. & Giannetti A., 2002 - Three-dimensional
burrow systems and taphofacies in shallowing-upward
parasequences, Lower Jurassic carbonate platform
(Calcari Grigi, Southern Alps, Italy) Facies, Erlangen,
47:57-82.
Monaco P., Giannetti A., Caracuel J. E. & Yébenes A.,
2005 - New shell-armoured burrows ( Ereipichnus
geladensis, n. ichnogenus) and associated echinoid
trace fossils from thè Prebetic of Alicante (Lower Cre-
taceous, SE Spain). Lethaia, Oslo, 38: 1-13.
Nickell L. A. & Atkinson R. J. A., 1995 - Functional
morphology of burrows and trophic modes of three
thalassinidean shrimp species, and a new approach to
thè classification of thalassinidean burrow morphol¬
ogy. Marine Ecology Progress Series, Halstenbek,
128: 181-197.
I
52
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Suchanek T. H., Colin P. L., McMurtry G. M. & Sucha-
nek C. S., 1986 - Bioturbation and redistribution of
sediment radionuclides in Enewetak Atoll lagoon
by callianassid shrimp; biological aspects. Bulletin
of Marine Science, Coral Gables (FL) 38 (1): 144-
154.
Vaugelas J. V. d., 1990 - Ecologie des callianasses
(Crustacea, Decapoda, Thalassinidea) et miléu recifal
Indo-Pacifique. Conséquences du remaniement sédi-
mentaire sur la distribution des materies humiques,
des métaux traces, et des radionucléides. Memoire
présenté à l’Université de Nice - Sophia Antipolis
pour l’obtention du Diplome d’Habilitation à diriger
des recherches en Sciences.
Vilas L., Arias C., Elizaga A., Garcìa de Domingo A. &
Lopez Olmedo F., 1982 - Consideraciones sobre el
Cretàcico inferior de la Zona de Jumilla-Yecla. Cuad-
ernos de Geologia Ibèrica, Madrid, 8: 635-650.
Winterer E. L. & Bosellini A., 1981 - Subsidence and
sedimentation on Jurassic passive Continental margin,
Southern Alps, Italy. AAPG Bulletin, Tulsa, 65 (3):
394-421.
Yébenes A., 1996 - Estratigrafia y estructura de la Serra
Gelada. Cuadernos de Geografia de la Universidad de
Valencia (60): 201-222.
Zempolich W. G., 1993 - The drowning succession in
Jurassic carbonates of thè Venetian Alps, Italy: a
record of supercontinent breakup, graduai eustatic
rise, and eutrophication of shallow-water environ-
ments. In Carbonate Sequence Stratigraphy - Recent
Developments and Applications. AAPG Memoir,
Tulsa, 57: 63-105.
Alice Giannetti - Geologisches Institut, Bonn Universitat, NuBallee 8, 53115 Bonn, Germany.
e-mail: giannetti@geo.uni-bonn.de
Paolo Monaco - Dipartimento Scienze della Terra, Università Perugia, piazza dell’Università, 06100 Perugia, Italy.
e-mail: pmonaco@unipg.it
Jesus E. Caracuel - Dpto. Ciencias de la Tierra y del Medio Ambiente, Universidad de Alicante, Apdo. 99, 03080 Alicante, Spain.
e-mail: jesus.caracuel@ua.es
Jesus M. Soria - Dpto. Ciencias de la Tierra y del Medio Ambiente, Universidad de Alicante, Apdo. 99, 03080 Alicante, Spain.
e-mail: jesus.soria@ua.es
Alfonso Yébenes - Dpto. Ciencias de la Tierra y del Medio Ambiente, Universidad de Alicante, Apdo. 99, 03080 Alicante, Spain.
e-mail: ayeb@telefonica.net
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Danièle Guinot, Antonio De Angeli & Alessandro Garassino
Discovery of thè oldest eubrachyuran crab from thè
Middle Jurassic (Bathonian) of Normandy (France)
The studied specimen was discovered in Normandy
(Calvados, France) in a quarry at Ranville, located near
thè city of Caen on thè right bank of Ome river. It was
collected from thè top of thè Hollandi Subzone ( Discus
Zone) of thè Calcaire de Langrune Fm., a limestone for-
mation of thè upper Bathonian (Middle Jurassic). The
Jurassic deposits at Ranville have long been a source of
interesting fossil materials (Rioult et al., 1991). The inter-
esting fauna from this region includes sponges, bryozo-
ans, gastropods, bivalves, ammonite and nautiloid cepha-
lopods, brachiopods, crinoids, and also several crustacean
families (Van Straelen, 1925), contrary to thè unfortunate
sentence in Guinot et al. (2005).
The studied specimen is an incomplete three-dimen-
sional, well-preserved carapace, ascribed to thè infraorder
Brachyura Latreille, 1802, and referred to thè family
Corystidae Samouelle, 1 8 19, in which it constitutes a new
genus and a new species (Guinot et al., work in progress).
The morphological characters of thè carapace exhibit
affinities with thè representatives of thè family Corysti¬
dae, especially Corystes Bosc, 1802, with C. cassive-
launus (Pennant, 1777) as type species. The similarities
are: carapace longer than broad and longitudinally ovate,
disposition of grooves on dorsal surface, front narrow and
with two teeth (or lobes), relatively wide orbits, raised
supra-orbital margin notched by two fissures, antero- and
postero-lateral margins not demarcated from each other,
postero-lateral margin hearing a subterminal tooth, and
posterior margin convex. The only differences are subtle
and concem thè proportions of thè body, slightly wider
anteriorly in thè studied specimen, thè distance between
thè two extraorbital teeth (fronto-orbital margin), more
important in thè studied specimen, thè tubercles of dorsal
surface, slightly stronger in thè studied specimen, thè
grooves, slightly more marked and smooth in thè stud¬
ied specimen (instead of faint and granulated in C. cas-
sivelaunus), and thè presence of small, smooth areas on
frontal and suborbitai regions in thè studied specimen
(instead of a dense and more uniform granulation of these
regions in C. cassivelaunus). Nevertheless, thè French
fossil corystid and C. cassivelaunus share thè same oma-
mentation of carapace, consisting of isolated tubercles in
thè anterior half and of clumps of 2-3 small granules in
thè posterior half.
Our knowledge of thè Mesozoic brachyurans is stili
limited because they are very rare in thè fossil record and
often are poorly preserved. The fossil Brachyura known
from thè Jurassic are represented by few primitive forms,
all belonging to thè Podotremata Guinot, 1977, namely
thè families Prosopidae v. Meyer, 1860, Homolodromii-
dae Alcock, 1900, Diaulacidae Wright & Collins, 1972,
Dynomenidae Ortmann, 1892, and Homolidae De Haan,
1839 (Glaessner, 1969; Guinot & Tavares, 2001; Sch-
weitzer et al., 2003; Schweitzer & Feldmann, 2005).
The family Prosopidae, recently revised by Wehner
(1988) and Muller et al. (2000), is an extinct family,
with many Mesozoic species, almost exclusively known
from carapace materials. Prosopidae, which is supposed
to have appeared in thè Lower Jurassic (Krobicki et al.,
2005), represents a case noted by Schweitzer & Feldmann
(2005) of Mesozoic genera disappearing during thè K/T
event, in contrast to others that became widespread during
thè Cenozoic. The fossil record suggests that thè family
Prosopidae is thè most ancient podotreme family. Only
one other family, thè Eocarcinidae Withers, 1932, if it is
truly brachyuran, is more ancient.
Forster (1979, 1985) regarded Eocarcinus praecur-
sor Withers, 1932, from thè Lower Jurassic (lower
Pliensbachian) of Great Britain, as thè putative ancestor
of thè Brachyura. He pointed out some morphological
characters similar to those of thè macruran Pseudopem-
phix Wiist, 1903, from thè Middle Triassic (Wellenkalk)
of Germany. Eoprosopon klugi Forster, 1986, from thè
Lower Jurassic (upper Pliensbachian) of Germany, was
considered an “intermediate form, connecting thè Pro¬
sopidae with thè oldest known crab-like Eocarcinus ” and
was included in thè Prosopidae (Forster, 1986). Later,
Guinot & Tavares (2001) discussed thè relationships
of Eocarcinus, stressing its morphological features. If
Eocarcinus Withers, 1932, is confirmed to be a brachy¬
uran crab with a paired spermatheca, it may deserve its
own superfamily, and thè Eocarcinidae would represent
thè oldest podotreme family.
The morphological characters of thè carapace of thè
French fossil corystid do not show evident correlations
with Pseudopemphix (Glypheoidea), nor with Eocarcinus
and Eoprosopon which have more backwardly enlarged
carapaces, a concave posterior margin, one developed
rostral spine, and complete postcervical and branchial
grooves. In thè same way, thè carapace of thè fossil corys¬
tid does not resemble that of thè Jurassic crabs of thè Pro¬
sopidae. Furthermore, thè fossil corystid does not belong
to other podotreme groups as Dromiacea, Homoloidea, or
Raninoidea.
Some representatives of thè Raninoidea, which
appeared at thè beginning of thè Cretaceous with three
genera ( Notopocorystes McCoy, 1849, Eucorystes Bell,
1863, and Cretacoranina Mertin, 1941), share some mor¬
phological characters of thè carapace with thè Jurassic
corystid, especially thè carapace that is elongate, more
enlarged forward, relatively convex longitudinally, and
thè fronto-orbital margin filling thè entire anterior margin.
We have excluded thè Jurassic corystid from thè Ranini-
dae for thè unbi lobate rostrum ( Notopocorystes excepted).
54
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
for thè preorbitai tooth not distinguished from thè rostral
spine, not raised and not delimited by a posterior groove.
Even though, Notopocorystes exhibits a bilobate rostrum,
as does thè Jurassic corystid, its rostral margins are con-
tinuous with thè preorbitai tooth that is not well raised and
not marked posteriorly. Moreover, thè trend of thè grooves
in thè anterior part of carapace in Notopocorystes (see
Withers, 1928; Wright & Collins, 1972; Collins, 1997;
Tucker, 1998; Schweitzer & Feldmann 2001) is different
from that of thè studied specimen, and Notopocorystes
generally shows a strong longitudinal median carina or
median row of tubercles, absent in thè studied specimen.
According to Wright & Collins (1972) Diaulax Bell,
1863, type genus of thè Diaulacidae, appeared in thè
Upper Jurassic and belongs to “a stock that diverged from
Dynomenidae already in thè Late Jurassic”. The Jurassic
fossil does not show any feature of a diaulacid crab.
As a result, thè Jurassic fossil is suggested to belong
to thè Eubrachyura and it is assigned to thè Corystidae.
Since thè studied specimen preserves only thè dorsal part
of carapace (ventral parts and pereiopods are lacking),
and in spite of an evident corystid facies, such an ascrip-
tion remains uncertain because thè Corystidae was known
only with three species from thè Miocene, Corystes
holsaticus (Noetling, 1881), C. latifrons (Lòrenthey in
Lòrenthey & Beurlen, 1929), and Gomezinus tuberculatus
Collins, Lee & Noad, 2003.
If our hypothesis that thè Jurassic specimen belongs
to thè Corystidae is correct, it represents thè oldest record
for thè Eubrachyura. Moreover, it allows development of
new hypotheses about thè probable evolutionary relation-
ships of brachyuran crabs. The hypothesis that thè clado¬
genesi Podotremata/Eubrachyura occurred prior to thè
Jurassic would be confirmed (Guinot & Tavares, 2001,
Collins J. S. H., 1997 - A systematic survey of thè genus
Notopocorystes McCoy, 1849 (Crustacea, Decapoda,
Raninidae). Bulletìn of thè Mizunami Fossil Museum,
Mizunami, 23, 1996 (1997): 75-87.
Forster R., 1 979 - Eocarcinus praecursor Withers (Deca¬
poda, Brachyura) from thè Lower Pliensbachian of
Yorkshire and thè early crabs. Neues Jahrbuch fur
Geologie und Palàontologie, Monatshefte, Stuttgart,
1979(1): 15-27.
Forster R., 1985 - Evolutionary trends and ecology of
Mesozoic decapod crustaceans. Transactions of thè
Royal Society of Edinburgh , Edinburgh, 76: 299-
304.
Glaessner M. F., 1969 - Crustacea Decapoda. In: Treatise
on Invertebrate Paleontology. Arthropoda 4 (2), Geo¬
logica! Society of America and University of Kansas,
Lawrence: R399-R533, R626-R628.
Guinot D. & Tavares M., 2001 - Une nouvelle famille
de Crabes du Crétacé, et la notion de Podotremata
Guinot, 1977 (Crustacea, Decapoda, Brachyura). Zoo-
systema, Paris, 23 (3): 507-546.
Guinot D., Wilson G. D. F. & Schram F. R., 2005 -
Jurassic isopod (Malacostraca: Peracarida) from Ran-
ville, Normandy, France. Journal of Paleontology,
Lawrence, 79 (5): 954-960.
Tab. 16). The studied specimen is most likely thè first
known Heterotremata.
The identities of thè earliest brachyurans are doubtful.
The putative Mississippian crab, Imocaris tuberculata
Schram & Mapes, 1984, is probably not a Brachyura
Dromiacea but revealed to be, after thè discovery of I.
colombiensis Racheboeuf & Villaroel, 2003 (Upper Car-
boniferous - Colombia), an Eocarida Brooks, 1962, and
a Pygocephalomorpha Beurlen, 1930 (Racheboeuf & Vil¬
laroel, 2003). But for thè time being such an assignment
is considered uncertain by Schram & Dixon (2004). The
Triassic species incertae sedis from New Mexico, Rioar-
ribia schrami Rinehart, Lucas & Heckert, 2003 (Rinehart
et al, 2003; Rinehart & Lucas, in press), supposed to be
an eubrachyuran crab, was stated by Schweitzer & Feld¬
mann (2005) not to be a decapod.
So, if our interpretation that thè carapace of thè French
corystid from thè Middle Jurassic of Normandy belongs
to thè Corystidae is correct, an older origination of most
basai Hetrotremata than supposed would be demonstrated.
Nevertheless, more evidence is required, especially ven¬
tral parts, to assert thè existence of thè Eubrachyura in
thè Jurassic. The origination of thè studied specimen and
of thè Heterotremata as a whole needs to be found in thè
more ancient fossil record.
The hypothesis that thè Hetrotremata probably evolved
in thè Jurassic Period is supported by thè presence of sev-
eral authentic heterotreme superfamilies/families in thè
fossil records known from thè Lower or thè Upper Cre-
taceous, including thè Dorippoidea MacLeay, 1838 (Vega
& Feldmann, 1992; Schweitzer & Feldmann, 2001), thè
Necrocarcinidae Forster, 1968, thè Carcineretidae Beur¬
len, 1930, and thè Hepatidae Stimpson, 1871 (Schweitzer
& Feldmann, 2000).
Krobicki M., Mtiller R & Zaton M., 2005 - Middle and
Upper Jurassic brachyuran crabs - phylogenetic and
palaeoenvironmental significance of their early evolu¬
tionary stage. Salt Lake City Annual Meeting (Octo-
ber 16-19, 2005), Geological Society of America,
Abstracts with Programs, 37 (7): 187.
Muller P., Krobicki M. & Wehner G., 2000 - Jurassic and
Cretaceous primitive crabs of thè family Prosopidae
(Decapoda: Brachyura) - their taxonomy, ecology and
biogeography. Annales Societatis Geologorum Polo-
niae, Warzawa, 70: 49-79.
Racheboeuf P. R. & Villaroel C., 2003 - Imocaris colom¬
biensis n. sp. (Crustacea: Decapoda) from thè Penn-
sylvanian of Columbia. Neues Jahrbuch fur Geologie
und Palàontologie Monatshefte, Stuttgart, (10): 577-
590.
Rinehart L. F., Lucas S. G. & Heckert A. B., 2003 - An
early eubrachyuran (Malacostraca: Decapoda) from
thè Upper Triassic Snyder Quarry, Petrified Forest,
north-central Mexico. In: Zeigler K.E., Heckert A.B.
& Lucas S.G. (eds), Paleontology of thè Snyder
Quarry Albuquerque, New Mexico Museum of Naturai
History & Science, Albuquerque, 24: 67-70.
Rinehart L. F. & Lucas S. G., in press - Reillustration of
Rioarribia schrami, an early eubrachyuran from thè
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
55
Upper Triassic petrified forest Formation of north-
central New Mexico. Bulletin of New Mexico Naturai
History Museum, New Mexico, 37.
Rioult M., Dugué O., Jan du Chéne R., Ponsot C., Fily G.,
Moron J.-M. & Vail R R., 1991 - Outcrop sequence
stratigraphy of thè Anglo-Paris basin, Middle to Upper
Jurassic (Normandy, Maine, Dorset). Bulletin Centres
de Recherche Exploitation-Production Elf Aquitaine,
15(1): 101-194.
Schram F. R. & Dixon C. J., 2004 - Decapod phylogeny:
addition of fossil evidence to a robust morphological
cladistic data set. Bulletin of thè Mizunami Fossil
Museum , Mizunami, 31: 1-19.
Schweitzer C. E. & Feldmann R. M., 2000 - New species
of calappid crabs from western North America and
reconsideration of thè Calappidae sensu lato. Journal
ofPaleontology, Lawrence, 74 (2): 230-246.
Schweitzer C. E. & Feldmann R. M., 2001 - New Cre-
taceous and Tertiary decapod crustaceans from west¬
ern North America. Bulletin of thè Mizunami Fossil
Museum, Mizunami, (28): 173-210.
Schweitzer C. E. & Feldmann R. M., 2005 - Decapod
crustaceans, thè K/T event, and Palaeocene recovery.
In: Koenemann S. & Jenner R. A. (eds.). Crustacea and
Arthropod Relationships. Taylor & Francis Group,
Boca Raton, London, New York, Singapore.
Schweitzer C. E., Feldmann R. M., Fam J., Hessin W.
A., Hetricks S. W., Nyborg T. G. & Ross R. M. L.,
2003 - Cretaceous and Eocene Decapod Crustaceans
from Southern Vancouver Island, British Columbia,
Canada. National Research Council Press, Ottawa.
Tucker A. B., 1998 - Systematics of thè Raninidae (Crus¬
tacea: Decapoda: Brachyura), with accounts of three
new genera and two new species. Proceedings of thè
Biological Society of Washington, Washington, 111
(2): 320-371.
Van Straelen V., 1925 - Contribution à l’étude des Crus-
tacés Décapodes de la période jurassique. Bulletin de
TAcadémie royale Belge, Bruxelles, Classe Sci., sér.
2, 7: 1-462.
Vega F. J. & Feldmann R. M., 1992 - Occurence of
Costacopluma (Decapoda: Brachyura: Retroplumi-
dae) in thè Maastrichtian of southern Mexico and its
paleobiogeographic implications. Annals of Carnegie
Museum, Pittsburgh, 61 (2): 133-152.
Wehner G., 1988 - Uber die Prosopiden (Crustacea, Deca¬
poda) des Jura. Unpublished degree thesis, Munchen.
Withers T. H. 1928 - New Cretaceous Crabs from Eng-
land and Syria. LIX. Annals and Magazine of Naturai
History, New York, ser. 10, 1: 456-460.
Wright C. W. & Collins J. S. H., 1972 - British Creta¬
ceous crabs. Palaeontographical Society Monograph,
London, 126: 1-114.
Fig. 1 - Corystid, n. gen., n. sp. from thè Middle Jurassic (Bathonian)
of Normandy (France) (Guinot et al., work in progress).
Danièle Guinot - Muséum national d’Histoire naturelle, Département Milieux et peuplements aquatiques, 61 rue Buffon,
F-75231 Paris cedex 05, France.
e-mail: guinot@mnhn.fr
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: agarassino@libero.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Oscar Hemàndez-Monzón, Francisco J. Vega & Marco A. Coutino
A review of Lophoranina cristaspina from thè Middle Eocene
of Chiapas, Mexico and evolutionary implications
A new locality found near thè Cerro de Mactumatza
(Fig. 1), has yielded more specimens with morphologi-
cal structures not described previously for Lophora¬
nina cristaspina Vega, Cosma, Coutino, Feldmann,
Nyborg, Schweitzer & Waugh, 2001, reported from
middle Eocene calcarenites of thè San Juan Formation,
that crops out near Tuxtla Gutiérrez, Chiapas (Fig. 2).
Differences in thè shape of lateral spines suggest a pos-
sible intraspecific variation, that may represent sexual
dimorphism. The review of morphology of more juve-
nile specimens suggests that generai shape of thè cara¬
pace and ornamentation if preserved through develop-
ment. The size range for this species is extended to
include specimens with relatively large carapaces.
A comparison of thè only Cretaceous species of this
genus and thè American and European forms suggest
that there were at least two lineages in thè evolution of
thè widespread Lophoranina.
Specimens are to be deposited in thè Museo de
Paleontologia "Eliseo Palacios Aguilera”, Instituto
de Historia Naturai y Ecologia de Chiapas, Tuxtla
Gutiérrez, Chiapas, under provisionai custody of F.
Vega.
Systematic paleontology
Order Decapoda Latreille, 1 802
Infraorder Brachyura Latreille, 1802
Section Podotremata Guinot, 1977
Subsection Archaeobrachyura Guinot, 1977
Superfamily Raninoidea De Haan, 1839
Family Raninidae De Haan, 1839
Genus Lophoranina Fabiani, 1910
Lophoranina cristaspina Vega, Cosma, Coutino,
Feldmann, Nyborg, Schweitzer & Waugh, 2001
Additions to description. Carapace subhexago-
nal, elongate (Fig. 3 - 1-15). Anterior margin slightly
concave, along with anterolateral spines represents
thè widest part of carapace; three anterolateral spines,
outermost either bifid or lobulate, middle one about as
long as outer one, but half its width, slightly concave;
inner spine semitriangular, as long as other two spines
but thè widest of all. Sternal segments fused into nearly
fiat piate, elements 1-3 subpentagonal, with spinose
anterolateral margins; element 4 with broad, concave
lateral margins. Carpus of cheliped subovai, with six
transverse terraces that lack spines; manus semirectan-
gular, with row of spines on lower margin that increase
in size toward distai part, where a sharp spine forms
highest part of cheliped.
Discussion. Higher taxonomic level are based on
Guinot & Quenette (2005). Morphologic variation
shown in some specimens of L. cristaspina is evident
in thè shape of thè external anterolateral spine. Most
specimens show a bifid spine, but others of thè same
size, have a lobulate shape of this spine. This morpho-
type is more scarce, and may reflect certain difference
related with sexual dimorphism, although this will
only be confìrmed when specimens with articulated
abdomina are found, as pointed out by Feldmann &
Schweitzer (in press).
Comparison with carapace shape and anterolateral
spines of L. precocious Feldmann, Vega, Tucker, Garcia-
Barrera & Avendano, 1996, from thè Maastrichtian
Angostura and Ocozocoautla formations near Tuxtla
Gutiérrez, Chiapas, suggest an evolutionary trend in
which thè two external spines of thè Cretaceous species
were fused into a bifid spine in thè Eocene species from
thè same region. The carapace with open lateral spines
and wider portion of carapaces at thè anterior margin is
also observed in L. bishopi Squires & Demetrion, 1992.
However, thè remaining European species (Via, 1959,
1965, 1969; Beschin et al., 2004), as well as L. geor¬
giana (Rathbun, 1935) from thè Oligocene of SE USA,
have a narrow anterior margin, with thè widest part of
thè carapace in thè anterior third of carapace length,
behind thè anterior margin. Based on these features,
thè following evolutionary trends may be suggested:
one includes thè Eocene American species, with broad
anterolateral spines; another lineage involves all thè
Tertiary European species, with relatively short ante¬
rior margins (Fig. 4). Presence of L. georgiana in thè
Oligocene of Georgia and Alabama may represent thè
influence of thè European linage, once thè American
lineage became extinct by late Eocene times.
60 meters
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
57
Fig. 1 - Location map of new outcrop near Cerro Mactumatza, south of
Tuxtla Gutiérrez, Chiapas.
Figs. 3 - 1-15) Lophoranina cristaspina from Cerro Mactumatza,
Tuxtla Gutiérrez, Chiapas. 1-2) Mirror images of specimens with bifìd
spinose and lobulate lateral spines, x 2.0. 3-4) Specimens with bifìd
spinose lateral spines, x 1.6. 5) Largest specimen, x 1.0. 6) Anterior
part of stemum, XI. 6. 7-8) Carpi of right and left chelipeds, x 2.8.
9) Outer view of right manus, x 3.0. 10) Inner view of left manus,
x 2.0. 11) Outer view of left manus, x 2.0. 12-13) Reconstruction of
specimens with bifìd and lobulate lateral spines. 14) Reconstruction of
stemum. 15) Reconstruction of left cheliped.
C
R AMERICA
E
T
A
C
E
O
u
s
E
o
c
E
N
E
O
L
I
G
O
C
E
N
E
EUROPA
L. straeloni
L. precocious
/
L. cristaspina L. bishopi
L. georgiana
Fig. 2 - Composite Cretaceous-Tertiary stratigraphic section at Cerro Fig. 4 - Possible evolutionary trends of Lophoranina, from thè Creta-
Mactumatza, including middle Eocene San Juan Formation. ceous species L. precocious.
58
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
References
Beschin C., Busulini A., De Angeli A. & Tessier G.,
2004 - The Eocene decapod crustacean fauna of thè
“Main” quarry in Arzignano (Vicenza-NE Italy) with
thè description of a new species of Raninidae. Lavori
- Società Veneziana di Scienze Naturali, Venezia, 29:
109-117.
Fabiani R., 1910-1 crostacei terziari del Vicentino. Bol-
letino del Museo Civico di Vicenza , Vicenza, 1: 1-40.
Feldmann R. M. & Schweitzer C., in press - Sexual
dimorphism in extinct and extant Raninidae (Deca-
poda: Brachyura). Annals of Carnegie Museum.
Feldmann R. M., Vega F. J., Tucker A. B., Garcia-Bar-
rera R & Avendano J., 1996 - The oldest record of
Lophoranina (Decapoda: Raninidae) from thè Late
Cretaceous of Chiapas, southeastem Mexico. Journal
of Paleontology, Lawrence, 70 (2): 296-303.
Guinot D., 1977 - Propositions pour une nouvelle classifi-
cation des Crustacés Decàpodes Brachyoures. Compte
Rendue Académie des Sciences de Paris , Serie D,
Paris, 285: 1049-1052.
Guinot D. & Quenette G., 2005 - The spermatheca in
podotreme crabs (Crustacea, Decapoda, Brachyura,
Podotremata) and its phylogenetic implications. Zoo-
systema, Paris, 27 (2): 267-342.
Haan W. de, 1833-1850 - Crustacea. In: Fauna Japonica
sive descriptio animalium, quae in itinere per Japo-
niam, jussu et auspiciis superiorum, qui summum in
India Baiava Imperium tenent, suscepto annis 1823-
1830, collegit, notis, observationibus et adumbratio-
nibus illustrativ. P. F. de Siebold & A. Amz (eds.).
London.
Latreille P. A., 1802-1803 - Histoire naturelle, général et
particuliére, des crustacés et des insectes. 3. F. Dufart ,
Paris.
Rathbun M. J., 1935 - Fossil Crustacea of thè Atlantic and
Gulf Coastal Plain. Geologica I Society of America,
Special Paper , Washington, 2.
Squires R. L. & Demetrion R. A., 1992 - Paleontology of
thè Eocene Bateque Formation, Baja California Sur,
Mexico. Naturai History Museum of Los Angeles County
Contributions in Science, Los Angeles, 434: 1-55.
Vega F. J., Cosma T., Coutino M. A., Feldmann R. M.,
Nyborg T. G., Schweitzer C. & Waugh D., 2001 - New
Middle Eocene Crabs (Crustacea: Decapoda) from
Chiapas, Mexico. Journal of Paleontology, Lawrence,
75 (5): 929-946.
Via L., 1959 - Decàpodos fósiles del Eoceno espanol.
Boletin Instituto Geològico y Minerò de Espana,
Madrid, 70: 331-402.
Via L., 1965 - Raninidos fósiles de Espana. Contribución
al conocimiento paleontològico de la familia
“Raninidae” (Crustàceos, Decàpodos). Boletin del
Instituto Geològico y Minerò de Espana, Madrid, 76:
233-275.
Via L., 1969 - Crustàceos decàpodos del Eoceno espanol.
Pirineos, Barcelona, 91-94: 1-479.
Oscar Hemàndez-Monzón - Facultad de Ciencias, UNAM, Ciudad Universitaria, Coyoacàn, México, D.F. 04510, Mexico.
e-mail: pipposkar@hotmail.com
Francisco J. Vega - Instituto de Geologia, UNAM, Ciudad Universitaria, Coyoacàn, México, D.F. 04510, Mexico.
e-mail: vegver@servidor.unam.mx
Marco A. Coutino - Museo de Paleontologia “Eliseo Palacios Aguilera”, Instituto de Historia Naturai y Ecologia de Chiapas,
Calzada de los Hombres Ilustres s/n, Parque Madero, Tuxtla Gutiérrez 29000, Chiapas, Mexico.
e-mail: coutinoj@hotmail.com
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Matùs Hyzny
Paleogene crab fauna of Borové Formation
V
(localities Durkovec and Hlinisko), western Carpathians, Slovakia
The Borové Formation is part of thè Podtatranskà
(Subtatric) Group, which is composed of sedimentary
rocks of Paleogene age. The Borové Formation is a typi-
cal transgressive formation with occurrences of marine
fauna, mostly bivalves and gastropods. The lithology
includes breccia, conglomerate, sandstone, limestone,
and rarely, also claystone. There are a few regional
lithostratigraphic units of thè Borové Formation in thè
Homàdska basin: thè Homàd Member, Chrast’ Member
and Tomàsovce Member (Gross et al., 1999). There are
some problems with thè stratigraphic dating of thè Borové
Formation because of no, or very few, important fossils
for stratigraphic biozonation. The age of each single
member of thè Borové Formation was determined mostly
indirectly. The Borové Formation has a rich macrofauna
with remains of decapod crustaceans, especially crabs
(Brachyura), which occur at several localities in thè
Homàdska basin.
The Tomàsovce Member represents thè uppermost
member of thè Borové Formation of thè Homàdska
and Sarisskà vrchovina uplands, dating to Priabonian -
Lower Oligocene (predominantly thè youngest Priabon¬
ian) time. The name of this member was taken from thè
village of Spisské Tomàsovce, and thè stratotype local-
ity is in thè Durkovec quarry. The Tomàsovce Member
represents a complex up to 150 m thick, composed of
altemating fine-grained sandstones and siltstones, with
pyrite concretions and scarce intercalations of medium-
grained carbonate arenites and fme-grained petromict
conglomerates. References to thè Tomàsovce Member
as thè sandstones with plants imprints and macrofauna“
have been known since thè 19th century (Hazslinszky,
1852; Miczynski, 1891; Staub, 1891). The Tomàsovce
Member has been thè object of paleontological study
by several authors (Gross et al., 1973; Filo & Sirànovà,
1996). The layers of thè Tomàsovce Member contain a
hydrophilous tropical flora (predominantly angiosperm
plants), a neritic macrofauna (predominantly bivalves)
and benthic foraminifers. The faunal assemblages are
characteristic of a neritic, marine environment (with thè
addition of a littoral thanatocoenosis) and a prevalence
of euryhaline forms.
Locality Durkovec is situated 1 km south from thè
village of Spisské Tomàsovce in thè Spisskà Novà Ves
district. The Durkovec quarry was open for thè extraction
of wall stone - sandstone. Extraction was performed in
thè past; thè Durkovec quarry was active about 150 years
ago. Modem extraction was opened on two levels. In thè
sandstones rich in bivalves and Paleogene flora some
specimens of decapod crustaceans also have been found,
specifically crabs. Crab remains of thè Durkovec local¬
ity were also known in thè past, but nobody studied this
assemblage. All of thè paleontological works and studies
of thè Paleogene sediments were focused mainly on mol-
luscs (Papsovà, 1970, 1973, 1975, 1977, 1978; Volfovà,
1960, 1963).
Locality Hlinisko is situated about 2.5 km south from
thè village Smizany in thè Spisskà Novà Ves district,
about 8 km east-south-east from thè Durkovec locality.
It is about 50 m long. Hlinisko is represented by layers
of sandstones and siltstones of thè Tomàsovce Member.
Pyrite concretions are very common. The rocks are
strongly eroded and irregularly disintegrated into layers
and locally disintegrated shale, unlike thè Durkovec
locality.
Both thè Durkovec and Hlinisko localities are very
rich in fossil remains, mainly subtropical flora and pelecy-
pods, but also including gastropods, echinoids, decapods,
and shark teeth. The Spisskà Novà Ves district has been
thè object of paleontological research by thè Civic
Museum in Spisskà Novà Ves since 1988 (Krempaskà,
1998). The first study about thè crab fauna from this area
was published by Hyzny (2006). Crab remains from thè
mentioned localities are well preserved. Rarely there were
found as complete, isolated specimens ( Ranina ( Ramina )
speciosa, Ranina sp.), so it is possible to describe thè
characters of thè pereiopods and claws in several cases.
Specimens of Ranina sp. were collected which probably
were preserved in living position, because thè orientation
of thè specimens is almost perpendicular to thè bedding
planes. At thè Durkovec quarry locality, there were also
collected some trace fossils referred to decapod crus¬
taceans: Thalassinoides igen. and Ophiomorpha igen.
A dose relationship between thè trace fossils and crab
specimens has not been determined until now.
Bittner (1875) mentioned an association of thè crab
Coeloma vigil and thè bivalve Pholadomya puschi from
thè marls of northem Italy. Fossils found at thè Durkovec
locality affimi this association. There are also some simi-
larities between thè crab fauna of thè Tomàsovce Member
and thè transitional layers of thè Molare Formation and
thè Rigoroso Mari in Italy (Allasinaz, 1987).
The environmental interpretation based on thè mac¬
rofauna and macroflora found at thè mentioned localities
for thè Spisskà Novà Ves district is assigned as a shallow
marine environment (maximum depth of 200 m).
The Decapoda species preliminarily identifìed from
deposits of thè Spisskà Novà Ves district include:
Raninidae: Ranina (Ranina) speciosa (Miinster, 1 840),
Ranina sp.
Calappidae: Calappilia sp.
Portunidae: IPortunites sp.
Geryonidae: Coeloma ( Coeloma ) vigil A. Milne
Edwards, 1865
60
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Fig. 1 - A) Ranina sp. Specimen orientateci almost perpendicular to thè bedding planes. Tomàsovce Member (Borové formation,
Priabonian, lower Oligocene), thè Durkovec locality, western Carpathians (Slovakia). B) Detail of thè same specimen. C) Ranina
sp. Ventral view on thè specimen with preserved claws. Tomàsovce Member, thè Durkovec locality. D) Coeloma ( Coeloma ) vigli A.
Milne Edwards, 1865. TomàSovce Member (Borové formation, Priabonian, lower Oligocene), thè Hlinisko locality, western Carpath¬
ians (Slovakia). E) Coeloma ( Coeloma ) vigil A. Milne Edwards, 1865. Tomàsovce Member, thè Durkovec locality. F) Calappilia sp.
Tomàsovce Member, thè Durkovec locality.
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
61
References
Allasinaz A., 1987 - Brachyura Decapoda oligocenici
(Rupeliano) del Bacino Ligure Piemontese. Bollettino
Museo Regionale Scienze Naturali Torino, Torino, 5
(2): 509-566.
Bittner A., 1875 - Die Brachyuren des vicentinischen
Tertiargebirges. Denkschriften der Kaiserlichen
Akademie der Wissenschaften, Mathematisch-
Natunvissenschaftliche Classe.
Filo I. & Sirànovà Z., 1996 - Tomàsovské vrstvy - nova
litostratigrafickà jednotka podtatranskej skupiny.
Geologické pràce, Geologickà sluzba Slovenskej
republiky, Bratislava, Spràvy 102: 41-49.
Gross P., Bucek S., Durkovic T., Filo I., Maglay J.,
Halouzka R., Karoli S., Nagy A., Spisàk Z., Zec
B., Vozàr J., Borza V., Lukàcik E., Janocko J.,
Jetel J., Kubes P., Kovàcik M., Zàkovà E., Mello
J., Polàk M., Sirànovà Z., Samuel O., Snopkovà
P., Rakovà J., Zlinskà A., Vozàrovà A., Zecovà K.,
1999 - Vysvetlivky ku geologickej mape Popradskej
kotliny, Homàdskej kotliny, levocskych vrchov,
Spissko-Sarisského medzihoria, Bachume a Sarisskej
vrchoviny. 1:50.000. Geologickà sluzba Slovenskej
republiky, Vydavatel’stvo Dionyza Stura, Bratislava.
Gross P., Papsovà J. & Kòhler E., 1973 - Sedimentologia
a stratigrafia vrchnej casti bazàlnej litofàcie centràl-
nokarpatského paleogénu od Sarisskej vrchoviny po
Liptovsku kotlinu. Manuskript - archlv Geologickà
sluzba Slovenskej republiky, Bratislava.
Hazslinszky F., 1852 - Das Thal der Schvinka bei Radàcs
im Sàroser Comitate, sudòstlich von Eperies. Jb. K.-
Kòn. geol. Reichsanst., Wien.
Hyzny M., 2006 - Kraby centràlnokarpatského paleogénu.
Bachelor thesis. Prirodovedeckà fakulta Univerzity
Komenského, Bratislava, 1-37.
1 Krempaskà Z., 1998 - Niektoré vysledky geologického
a paleontologického vyskumu v lome Durkovec v
Slovenskom raji. Natura Carpatica, 39: 27-32.
Miczynski K., 1891 - Uber einige Pflanzreste von Radacs
bei Eperies, Comitat sàros. Mitt. Jb. Ung. geol. Anst.,
Budapest, 9.
Staub M., 1891 - Etwas iiber die Pflanzen von Radàcs bei
Eperies. Mitt. Jb. Ung. geol. Anst., Budapest, 9.
Papsovà J., 1970 - Zàverecnà spràva o makrofaune
paleogénu Liptovskej kotliny na liste Ruzomberok
/1:50.000/. Ciastkovà zàverecnà spràva za rok 1970.
Manuskript - archlv Geologicky ustav Dionyza Stura,
Bratislava.
Papsovà J., 1973 - Sedimentologia a stratigrafia vrchnej
casti bazàlnej litofàcie centràlnokarpatského paleo¬
génu od Sarisskej vrchoviny po Liptovsku kotlinu.
Dielcia zàverecnà spràva za rok 1973. Manuskript
- archlv Geologicky ustav Dionyza Stura, Bratislava.
Papsovà J., 1975 - Makrofauna strednej casti Liptovs¬
kej kotliny na listoch Huty, Demanovà, Smrecany,
Liptovsky Hràdok. Ciastkovà spràva za rok 1975.
Manuskript - archlv Geologicky ustav Dionyza Stura,
Bratislava.
Papsovà J., 1977 - Celkové zhodnotenie makrospo-
locenstiev Zàpadnych Karpàt. Ciastkovà zàverecnà
spràva za rok 1977. Manuskript - archlv Geologicky
ustav Dionyza Stura, Bratislava.
Papsovà J., 1978 - Makrofauna paleogénu vychodnej
casti Liptovskej kotliny. Ciastkovà spràva za rok
1978. Manuskript - archlv Geologicky ustav Dionysa
Stura, Bratislava.
Staub M., 1891 - Etwas uber die Pflanzen von Radàcs bei
Eperies. Mitt. Jb. Ung. geol. Anst., Budapest, 9.
Volfovà J., 1960 - Stratigraficko-ekologické vyhodno-
tenie makrofaunistickych lokalit centràlneho pod-
tatranského paleogénu /List Vysoké Tatry - Strba/.
Manuskript - archlv Geofond, Bratislava.
Volfovà J., 1963 - Zàverecnà spràva o makrofaune na
liste Hranovnica /L50.000/. Manuskript - archlv
Geologicky ustav Dionyza Stura, Bratislava.
il
Matùs Hyzny - Department of Geology and Paleontology, Faculty of Naturai Sciences, Comenius University in Bratislava,
Mlynskà Dolina G, 842 1 5 Bratislava, Slovakia.
e-mail: hyzny.matus@gmail.com
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Hiroaki Karasawa & Hisayoshi Kato
New prosopid crabs (Crustacea, Decapoda, Brachyura) from thè
Upper Jurassic Torinosu Group, Shikoku, Japan
The extinct podotreme family Prosopidae v. Meyer,
1860, is well known from thè Jurassic and Cretaceous
deposits of thè Tethys realm (Miiller et al., 2000). Previ-
ously known prosopids from thè north Pacific have been
Pithonoton inflatum Collins & Karasawa, 1993, from
thè Upper Cretaceous of Japan, Nipponopon hasegawai
Karasawa, Kato & Terabe, 2006, from thè Lower Creta¬
ceous (Barremian) of Japan, and Nodoprospon sp. from
thè Upper Jurassic (Oxfordian) of Japan (Collins & Kara¬
sawa, 1993; Karasawa et al., 2006; Kato et al., 2007).
The purpose of thè present study is to describe new
prosopid crabs from thè Upper Jurassic Torinosu Group
of Shikoku, Japan. The specimens were collected from
shale of thè Yatsuji Formation (Kimura, 1956) of thè
Torinosu Group exposed at Ogawa, Sakawa-cho, Kochi
Prefecture (33°29'13"N; 133°14'28"E). Crabs occurred
in deposits, in association with mollusks, Parallelodon
(Torinosucatella) kobayashii (Tamura), Entolium yat-
sujiense Kurata & Kimura, Chlamys iboibo Kurata &
Kimura, Aequipecten ? spp., Parvamussium habunoka-
xvense (Kimura), and Goniomya ( Goniomya ) nonvscripta
Tamura (K. Mimoto & T. Hirota, per. com.). Shiraishi et
al. (2005) showed that thè strontium isotopie age of thè
Yatsuji Formation is 146.1-148.4 Ma (middle Tithonian).
The described and figured specimens are housed in thè
Mizunami Fossil Museum (MFM) and thè Sakawa Geol-
ogy Museum (SGM), Sakawa-cho, Kochi Prefecture,
Japan.
Systematic paleontology
Family Prosopidae v. Meyer, 1860
Subfamily Goniodromitinae Beurlen, 1932
Genus Goniodromites Reuss, 1 859
Goniodromites hirotai sp. nov.
Diagnosis: large sized Goniodromites. Carapace
pentagonal, much wider than long, widest at about mid-
length. Upper orbitai margin rimmed with broadly trian-
gular outer orbitai tooth. Anterolateral margin sinuous,
sharp, with minute spine just in front of cervical notch and
small epibranchial spine. Posterolateral margin sinuous;
sharp, tumid margin continuous with anterolateral margin
being within anterior third. Dorsal surface irregularly
tuberculate, slightly convex transversely and longitudi-
nally. Mesogastric region barely differentiated from pro-
togastric regions, with weak, shallow median depression.
Cervical groove well defìned. Branchiocardiac grooves
poorly defìned.
Etymology: thè specific name honors Mr. T. Hirota
who first discovered thè Jurassic decapod fauna from
Shikoku.
Material examined: MFM247,020 (holotype) and
SGM 1297 (paratype) collected by T. Hirota.
Description. Carapace large, pentagonal in outline,
wider than long, length about 82% carapace width,
widest at about mid-length. Front-orbital margin about
82% maximum width. Frontal margin narrow, about
25% maximum width, with shallow median notch. Upper
orbitai margin continuous with frontal margin, long,
gently convex, rimmed, without fissures. Outer orbitai
tooth broadly triangular, directed anterolaterally. Ante¬
rolateral margin sinuous, sharp, short, hearing minute
spine just in front of cervical notch; epibranchial spine
present, small, slightly directed backward. Posterolateral
margin sinuous, tapering posteriorly; sharp, tumid margin
continuous with anterolateral margin reaching at anterior
third. Posterior margin slightly concave, rimmed, about
42% maximum width. Dorsal surface irregularly tuber¬
culate on outer mould but nearly smooth on internai
mould, slightly convex transversely and longitudinally,
with poorly defìned regions. Front downtumed, sulcate.
Epigastrio regions elevated. Mesogastric region pyriform,
maximum width about 32% carapace width, barely dif¬
ferentiated from protogastric regions, with weak, shal¬
low median depression; anterior mesogastric process
separated from epigastrio regions by rather deep grooves.
Hepatic and protogastric regions not differentiated.
Cervical groove well defìned, sinuous; lateral elements
nearly straight; axial element concave. Urogastric region
slightly vaulted longitudinally, separated from branchial
regions by shallow, posteriorly converged grooves. Car-
diac region triangular, broadest anteriorly, defìned by
broad, shallow grooves. Branchiocardiac grooves poorly
defìned, lateral elements nearly parallel lateral elements
of cervical groove. Meso- and metabranchial regions not
differentiated. Intestinal region weakly depressed.
Discussion. Since Glaessner ( 1 929) has treated Gonio¬
dromites as thè subgenus of Pithonoton v. Meyer, 1842,
subsequent workers used Goniodromites as thè subgenus
of Pithonoton and/or synonymized Goniodromites with
Pithonoton. Most recently, Feldmann et al. (2006) recog-
nized Goniodromites as a separate genus and included 14
species in it. We agree with their opinion on thè generic
status of Goniodromites. The present species appears
to possess internai dorsal carapace characters most like
those of Goniodromites etalloni (Gemmellaro, 1 869) from
thè Tithonian of Sicily and Romania (Gemmellaro, 1869;
3™ SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
63
Patrulius, 1966). However, thè irregularly tuberculated
dorsal carapace readily distinguishes G. hirotai from G.
etaìloni. The present species bears an anterolateral spine
anterior to thè cervical notch and one epibranchial spine,
but G. etaìloni lacks anterolateral spines.
De Angeli & Garassino (2006) synonymized G. etal-
loni with Goniodromites bidentatum Reuss, 1859, from
thè Tithonian of Europe, whereas Feldmann et al. (2006)
recognized G. etaìloni as a distinct species. According to
Feldmann et al. (2006), we treat G. etaìloni as a separate
status because in G. etaìloni thè carapace has a smooth
upper orbitai margin, no anterolateral tooth, and a poste-
rior mesogastric region with a weak median depression
and thè maximum carapace width is at about mid-length.
Although hitherto known members of Goniodromites
have been known from thè Middle Jurassic (Bajocian)-
Upper Cretaceous (Cenomanian) of thè Tethys realm (Feld¬
mann et al., 2006), G. hirotai sp. nov. and G. sakawense sp.
nov. represent thè first record from thè north Pacific realm.
Goniodromites sakawense sp. nov.
Diagnosis: small sized Goniodromites. Carapace
pentagonal, slightly wider than long, widest at outer
orbitai angle. Frontal margin about 40% maximum width,
composed of two broadly triangular teeth. Upper orbitai
margin nearly straight, rimmed, fringed with minute ser-
rations. Lateral margins nearly straight without serrations
and spines. Posterior margin concave, rimmed; width
about 50% maximum width. Dorsal surface densely
rugose, gently convex transversely and longitudinally.
Epigastrio regions well defined, transversely ridged.
Mesogastric region without posterior median depression.
Cervical groove well defined. Urogastric region separated
from cardiac region by shallow, gently concave groove.
Branchiocardiac grooves well developed.
Etymology: thè trivial name notes thè occurrence of
thè specimens in Sakawa-cho.
Material examined: MFM247,021 (holotype) and
SGM1298 (paratype) collected by T. Hirota.
Description. Carapace small, pentagonal in outline,
length about 90% maximum carapace width, widest at
outer orbitai angle. Front-orbital margin slightly shorter
than maximum width. Frontal margin about 40% maxi¬
mum width, composed of two broadly triangular teeth,
medially notched, United with upper orbitai margin.
Upper orbitai margin long, nearly straight, rimmed,
fringed with minute serrations; outer orbitai angle small,
triangular, directed anterolaterally. Lateral margins nearly
straight, tapered posteriorly, with cervical and branchio¬
cardiac incisions. Posterior margin concave, rimmed;
width about 50% maximum width. Dorsal surface
densely rugose, gently convex transversely and longi¬
tudinally. Front downtumed, sulcate. Epigastrio regions
well defined, transversely ridged. Mesogastric region
pyriform, maximum width about 36% carapace width,
barely differentiated from protogastric regions; anterior
mesogastric process separated from epigastrio regions by
deep grooves. Hepatic and protogastric regions not dif¬
ferentiated. Cervical groove well defined, sinuous; lateral
elements gently convex or nearly straight; axial element
concave. Urogastric region separated from cardiac region
by shallow, gently concave groove. Cardiac region trian¬
gular, broadest anteriorly. Intestinal region long, defined
by shallow grooves joining branchiocardiac grooves.
Branchiocardiac grooves well developed; lateral elements
nearly parallel lateral elements of cervical groove. Meso-
and metabranchial regions not differentiated.
Discussion. The present species is similar to Gonio¬
dromites serratum Beurlen, 1929, from thè Oxfordian-
Tithonian of Germany, Belgium, and Poland (Feldmann
et al., 2006) and Goniodromites sp. Form A from thè
Oxfordian of Romania (Feldmann et al., 2006). This spe¬
cies differs from G. serratum and G. sp. Form A in that thè
lateral margins are smooth without serrations and spines,
and thè carapace is slightly wider than long. A narrow
posterior mesogastric region in G. sakawense does not
bear a longitudinal groove, while G. serratum has a wide
posterior mesogastric region with a weak median groove.
Genus Pithonoton v. Meyer, 1 842
Pithonoton iyonofutanajima sp. nov.
Diagnosis: small to moderate sized Pithonoton Cara¬
pace elongate-pentagonal, much longer than wide, widest
at about outer orbitai angle. Frontal margin rimmed, about
40% maximum width, composed of two rounded lobes,
weakly notched medially. Upper orbitai margin slightly
concave, rimmed. Lateral margins slightly sinuous, with
cervical incision. Posterior margin concave, rimmed;
width about 45% maximum width. Dorsal surface
densely rugose, gently convex transversely and longitu¬
dinally. Front strongly protruded anteriorly. Epigastric
regions weakly ridged transversely. Cervical groove well
defined. Branchiocardiac grooves much shallower cervi¬
cal groove.
Etymology: thè specific name derived from “Iyonofu-
tanajima” meaning Shikoku in thè Japanese mythological
age.
Material examined: MFM247,022 (holotype) and
SGM1299 (paratype) collected by T. Hirota.
Description. Carapace small to moderate, elon¬
gate-pentagonal in outline, width about 87% maximum
carapace width, widest at about outer orbitai angle.
Front-orbital margin slightly shorter than maximum
width. Frontal margin rimmed, about 40% maximum
width, composed of two rounded lobes, weakly notched
medially, united with upper orbitai margin. Upper orbitai
margin long, slightly concave, rimmed; outer orbitai
angle small, triangular, directed anterolaterally. Lateral
margins slightly sinuous, tapered posteriorly, with cervi¬
cal incision. Posterior margin concave, rimmed; width
about 45% maximum width. Dorsal surface densely
rugose, gently convex transversely and longitudinally.
Front downtumed, strongly protruded anteriorly, sulcate.
Epigastric regions weakly ridged transversely. Mes¬
ogastric region pyriform, maximum width about 33%
carapace width, barely differentiated from protogastric
regions; anterior mesogastric process separated from
epigastric regions by rather deep grooves. Hepatic and
protogastric regions not differentiated. Cervical groove
well defined, sinuous; lateral elements nearly straight;
axial element concave. Urogastric region slightly vaulted
longitudinally. Cardiac region triangular, broadest anteri¬
orly, defined by broad, shallow grooves. Branchiocardiac
64
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Figs. 1-7 - 1) Nodoprosopon sp., SGM1300. a) Silicon cast of extemal mould of carapace; b) internai mould of carapace; c) extemal
mould of carapace. 2) Pithonoton iyonofutcmajima sp. nov., MFM247,022 (holotype), dorsal view of carapace. 3) Pithonoton iyonofu-
tanajima sp. nov., SGM1299 (paratype), dorsal view of carapace. 4) Goniodromites hirotai sp. nov., SGM1297 (paratype), dorsal view
of carapace. 5) Goniodromites sakawense sp. nov., MFM247,02 1 (holotype), dorsal view of carapace. 6) Goniodromites sakawense
sp. nov., SGM1298 (paratype), dorsal view of carapace. 7) Goniodromites hirotai sp. nov., MFM247,020 (holotype), dorsal view; a)
Silicon cast of extemal mould of carapace; b) internai mould of carapace, la-c, scale bars = 2 mm; 2, 3, 5, 6, scale bars = 5 mm; 4,
7a-b, scale bars = 10 mm.
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
65
grooves shallow, lateral elements nearly parallel lateral
elements of cervical groove. Meso- and metabranchial
regions not differentiated.
Discussion. The present new species resembles
Pithonoton marginatum (v. Meyer, 1842) from thè Kim-
meridgian-Tithonian of Europe (Garassino et al., 2005) and
Pithonoton elongatum (v. Meyer, 1 860) from thè Middle-
Upper Jurassic of Germany. The elongated front with two
rounded lobes, weakly raised epigastric regions, and weak
branchiocardiac grooves readily distinguish P iyonofu-
tanajima from P. marginatum. This species differs from P
elongatum in having a wide front with two rounded lobes, a
wide posterior margin, and weak branchiocardiac grooves.
The carapace in P. elongatum is much longer than wide.
Pithonoton iyonofutanajima represents thè second
record of Pithonoton from thè north Pacific rim. Recogni-
tion of thè present species extends thè geologie range for
1
Collins J. S. H. & Karasawa H., 1993 - The Cretaceous
crab, Pithonoton inflatum from Hokkaido, Japan. Sci¬
ence Reports of thè Toyohashi Museum of Naturai
History, Toyohashi, 3: 17-20.
De Angeli A. & Garassino A., 2006 - Catalog and bibli-
ography of thè fossil Stomatopoda and Decapoda from
Italy. Memorie della Società italiana di Scienze natu¬
rale e del Museo civico di Storia naturale di Milano,
Milano, 35 (1): 3-96.
Feldmann R. M., Lazar I. & Schweitzer C. E., 2006
-New crabs (Decapoda: Brachyura: Prosopidae) from
Jurassic (Oxfordian) sponge bioherms of Dobrogea,
Romania. Bulletin of thè Mizunami Fossil Museum,
Mizunami, 33: 1-20.
Garassino A., De Angeli A. & Schweigert G., 2005 -
Brachyurans from thè Upper Jurassic (Kimmeridgian-
Tithonian) of Pfalzpaint and Breitenhill (Bavaria, S
Germany). Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano,
Milano, 146 (1): 69-78.
Gemmellaro G. G., 1869 - Crostacei. In: Studi paleon¬
tologici sulla fauna del calzare a Terebratula janitor
del Nord di Sicilia. Stabilimento Tipografico Lao,
Palermo, Parte I: 11-18.
Glaessner M. F., 1929 - Crustacea Decapoda. In: Fos-
silium Catalogus: Ammalia. J. F. Pompeckj (ed.). W.
Junk, Berlin, 41: 1-464.
Karasawa H., Kato H. & Terabe K., 2006 - A new
member of thè family Prosopidae (Crustacea: Deca¬
poda: Brachyura) from thè Lower Cretaceous of
thè genus known from thè north Pacific rim back to thè
Tithonian.
Subfamily Prosopinae v. Meyer, 1 860
Genus Nodoprosopon Beurlen, 1928
Nodoprosopon sp.
Material examined: SGM1300 collected by T.
Hirota.
Discussion. The Japanese species closely resembles
Nodoprosopon heydnei (v. Meyer, 1860) from thè Oxford-
ian-Tithonian of Europe (Kato et al., 2007). Other mate¬
rial is reported from thè Oxfordian Somanakamura Group
of Fukushima Prefecture, Japan (Kato et al., 2007).
Japan. Revista Mexicana de Ciencias Geológicas,
Mexico, 23: 344-349.
Kato H., Takahashi T. & Taira M., 2007 - Decapod crus-
taceans from thè upper Jurassic of Somanakamura
Group, northeast Japan: first record of thè Jurassic
crab from thè circum-Pacifìc region. Abstracts with
Programs, The 156 Regalar Meeting of thè Palaeon-
tological Society of Japan, Takushima.
Kimura T., 1956 - The Torinosu Group and thè Tori-
nosu Limestone in thè Togano and Go Basins, Kochi
Prefecture. The Journal of thè Geological Society of
Japan, Tokyo, 62: 515-526.
Miiller P., Krobicki M. & Wehner G., 2000 - Jurassic and
Cretaceous primitive crabs of thè family Prosopidae
(Decapoda: Brachyura) - their taxonomy, ecology and
biogeography. Annales Societatis Geologorum Polo-
niae, Kraków, 70: 49-79.
Patrulius D., 1966 - Les Décapodes du Tithonique
inférieur de Wozniki (Carpates Polonaises Occiden-
tales). Annales de la Société Géologique de Pologne,
Warzawa, 36 (4): 495-517.
Reuss A. E., 1 859 - Zur Kenntnis fossiler Krabben. Denk-
schriften der kaiserlichen Akademie der Wissenschaf-
ten in Wien, Wien, 17: 1-90.
Shiraishi S., Hayasaka Y., Takahashi Y., Tanimizu M.,
Ishikawa T., Matsuoka J., Murayama M. & Kano
A., 2005 - Strontium isotopie age of thè Torinosu
Limestone in Niyodo Village, Kochi Prefecture, SW
Japan. The Journal of thè Geological Society of Japan,
Tokyo, 111: 610-623.
Hiroaki Karasawa - Mizunami Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifii 509-6132, Japan.
e-mail: GHA06103@nifty.com
Hisayoshi Kato - Naturai History Museum and Institute, Chiba, Aoba-cho, Chiba 260-8682, Japan.
e-mail: katoh@chiba-muse.or.jp
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Michal Krobicki, Pài Miiller & Michal Zatori
Middle and Upper Jurassic European prosopid crabs,
phylogeny and palaeoenvironments
The most primitive crabs (Brachyura) of thè extinct
family Prosopidae are represented mostly by Jurassic
and Cretaceous species. The origin of these crabs is more
or less enigmatic and mainly based on morphological
changes in thè surface of their carapaces. Fossil evidence
suggests that thè family Prosopidae is ancestral to all
other brachyurans including Dromiacea and Eubrachyura,
with very dose phylogenetic relations to Homolodromi-
idae, Dromiidae, Homolidae, Latreillidae, Dynomenidae,
Xanthidae, Cyclodorippoidea and Calappoidea. Prosop-
ids appeared in thè late Early Jurassic (Pliensbachian)
and therefore most probably accommodate thè ancestors
of all brachyurans (Miiller et al., 2000). Forster (1979),
who described thè oldest crab species - Eocarcinus prae-
cursor Withers, 1932, suggested that this form is in many
respects transitional between thè macruran Glypheoidea
and thè early brachyurans. This idea is widely accepted
suggesting that this early Pliensbachian Eocarcinus is a
transitional form between thè Triassic glypheoids ( Pseu -
dopemphix) and earliest, Late Pliensbachian prosopids
( Eoprosopon klugi Forster). Meyer (1860) was thè fìrst to
report prosopid crabs and recognised their dromiid affìni-
ties. He arranged thè 25 known species into a new family
Prosopidae. However, thè very dose relationship of pro¬
sopids and thè Homolodromiidae is universally accepted
(see e.g., Wehner, 1988; Schweitzer et al., 2004), and
seems to be well established by thè high degree of simi-
larity in carapace traits, pattems of furrows, ventral parts,
and, partly, chelae as well.
Most prosopid species were found in thè Upper Juras¬
sic limestones, while thè Early and Middle Jurassic fossil
record of crabs is very poor (e.g., Forster, 1985; Wehner,
1988; Miiller et al., 2000; Schweigert, 2006). Prosopids
had their climax during this time with a wide distribution
in sponge-microbial (Oxfordian) buildups and/or coral
reef (Kimmeridgian-Tithonian) environments of Europe
(Pithonoton, Coelopus, Longodromites , Prosopon, Foers-
teria, Nodoprosopon, Lecythocaris, Glaessneropsis).
Early-Middle Jurassic prosopid crabs, and therefore,
presumably also brachyuran evolution, started on shal-
low-sea, soft bottom environments, where their habitat
was probably closely related to oolitic and coral facies,
crinoid and also molluscan buildups/shell accumulations,
and moderatly deep grey clays sedimentation of Bajo-
cian-Bathonian age (Krobicki et al., 2005). Soon after,
thè world-wide expansion of epicontinental seas, associ-
ated with thè Callovian transgression, allowed thè forma-
tion of bioherms and reefs in thè Late Jurassic, creating
ideal ecological niches for thè rapidly evolving prosopids
(Forster, 1985). Numerous crab species following thè
Oxfordian growth of organogenie, mainly sponge-bearing
buildups, documented ecological (probably symbiotic)
relationships between thè faunal elements (e.g., Miiller
et al., 2000). These Late Jurassic cyanobacterial-sponge
limestones were deposited in deep sublittoral to littoral
settings, containing abundant sponges and sporadically
hermatypic corals as well (Matyszkiewicz et al., 2006
with references cited therein).
The Mid- to Late Jurassic wide distribution of shallow
water environments (maximum a few hundred metres)
and in particular thè varied reefal habitats around thè
Tethys Ocean, contributed much to thè rapid diversifica-
tion and success of early crabs (Forster, 1985; Miiller et
al, 2000). The early episode of crab evolution has been
connected with their climax during Late Jurassic time
with a wide distribution, probably developed under tropi¬
cal and subtropical conditions in shallow, warm waters
of thè sponge and sponge-coral reefs in thè Oxfordian-
Kimmeridgian, as well as coral reefs in thè Tithonian
(thè so-called Stramberk-type limestones) of thè northem
Tethyan Ocean. The isolated occurrences of Late Jurassic
prosopids outside thè centres of “reef ’ sedimentation sug-
gest that this fauna adapted poorly to non-reefal palaeo-
ecological conditions (Miiller et al., 2000). After retreat
of reefal facies at thè Jurassic-Cretaceous boundary, crab
development decreased and, therefore, thè Cretaceous
fossil record of this fauna is poor and spatially dispersed.
Their closely related descendants, thè homolodromi-
ids, preferentially inhabited soft muddy bottoms in
deeper, colder waters. Such ecological displacement to
deeper habitats is well documented by Tertiary fossils
(cf. Feldmann et al., 1991; Collins, 1997). Some homolo-
dromiids actually live even deeper than 1.000 m (Guinot
& Forges, 1981).
References
Collins J. S. H., 1997 - Fossil Homolidae (Crustacea;
Decapoda). Bulletin of thè Mizunami Fossil Museum ,
Mizunami, 24: 51-71.
Feldmann R. M., Tucker A. & Berglund R. E., 1 99 1 —
Fossil crustaceans. National Geographical Research
and Exploration, Washington, 7: 352-363.
Forster R., 1979 - Eocarcinus praecursor Withers
(Decapoda, Brachyura) from thè Lower Pliensbachian
of Yorkshire and thè early crabs. Neues Jahrbuch fìir
Geologie und Palàontologie, Monatshefte, Stuttgart,
1: 15-27.
Forster R., 1985 - Evolutionary trends and ecology of
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
67
Mesozoic decapod crustaceans. Transactions of thè
Royal Society of Edinburgh, Earth Sciences , Edin¬
burgh, 76: 299-304.
Guinot D. & de Forges B. R., 1981 - Homolidae, rares ou
nouveaux, de l’Indo-Pacifique (Crustacea, Decapoda,
Brachyura). Bulletin du Muséum national d’Histoire
naturelle, Paris, 4 ser., 3, sec. A, 2: 523-581.
Krobicki M., Miiller P. & Zaton M., 2005 - Middle and
Upper Jurassic brachyuran crabs - phylogenetic and
palaeoenvironmental significance of their early evolu-
tionary stage. The Geologica l Society of America, GSA
Annual Meeting & Exposition , Salt Lake City, 16-19
October 2005, Abstracts and Programs: 187.
Matyszkiewicz J., Krajewski M. & Zaba J., 2006 - Struc-
tural control on thè distribution of Upper Jurassic car¬
bonate buildups in thè Kraków - Wielun Upland (south
Poland). Neues Jahrbuch fur Geologie und Palàonto-
logie, Monathshefte , Stuttgart, 3: 182-192.
Meyer H. von, 1860 - Die Prosoponiden oder die Fami-
lie der Maskenkrebse. Palaeontographica, Kassel, 7:
183-222.
Miiller P., Krobicki M. & Wehner G., 2000 - Jurassic and
Cretaceous primitive crabs of thè family Prosopidae
(Decapoda: Brachyura) - their taxonomy, ecology
and biogeography. Annales Societatis Geologorum
Poloniae, Kraków, 70: 49-79.
Schweigert G., 2006 - A specimen of Prosopon hebes v.
Meyer, 1840 (Decapoda: Brachyura: Prosopidae) from
thè Middle Jurassic of SW Germany. Neues Jahrbuch
fur Geologie und Palàontologie, Monathshefte, Stutt¬
gart, 6: 361-370.
Schweitzer C. E., Nyborg T. G. & Feldmann R. M. &
Ross R. F. M., 2004 - Homolidae de Haan, 1839 and
Homolodromiidae Alcock, 1900 (Crustacea: Deca¬
poda: Brachyura) from thè Pacific Northwest of North
America and a reassessment of their fossil records.
Journal ofPaleontology, Fawrence, 78 (1): 133-149.
Wehner G., 1988 - Ùber die Prosopiden (Crustacea,
Decapoda) des Jura. Inaugunal - Dissertation zur
Erlangung des Doktorgrades der Fakultàt fur Geo-
wissenschaften der Ludwig-Maximilians-Universitàt
zu Miinchen.
This work was financially supported by thè grants from thè Agh University of Science and Technology in Kraków (MK).
Michal Krobicki - AGH University of Science and Technology, Mickiewicza 30, 30-059 Kraków, Poland.
e-mail: krobicki@geol.agh.edu.pl
Pài Miiller - Geological Institute of Hungary, Fòldtani Intézet, Stefania ut. 14, 1143 Budapest, Hungary.
e-mail: mullerp@mafi.hu
Michal Zaton - Faculty of Earth Sciences, B^dzinska 60, 41-200 Sosnowiec, Poland.
e-mail: mzaton@ultra.cto.us.edu.pl
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Cristiano Larghi & Andrea Tintori
First record of a decapod from thè Meride Limestone:
new data from one of thè best Ladinian (Middle Triassic)
taphonomic Windows of a transitional environment
The Middle Triassic Italian and Swiss localities of
Besano-Viggiù (Lombardy) and Monte S. Giorgio-Meride
(Canton Ticino) are sites where Triassic marine deposits
have been studied through continuous scientifìc excava-
tions over a period of more than 150 years. The Monte
San Giorgio area was included in thè World Heritage
List of UNESCO in July, 2003, and thè Italian authori-
ties are actually working to extend thè area to thè Ital-
ian sites belonging to thè same formations. The Triassic
sequence is composed of distinct stratigraphical units, thè
most fossiliferous of which are thè upper Anisian-lower
Ladinian Besano Formation (or Grenzbitumenzone), and
thè Ladinian (Middle Triassic) Meride Limestone, thè
upper levels of which are named Kalkschieferzone by thè
Swiss authors (Senn, 1924). The sequence records life in
a tropical lagoon, sometimes stagnant at thè bottom, and
hostile to thè benthos. The Kalkschieferzone in particular
is indicative of a hot climate, subject to seasonal drought
with conditions between fully marine or hypersaline
and sometimes ffesh water; its soft-body fossil biota is
thè result of a typical taphonomic window of a transi¬
tional environment (cfr. Briggs & Gali, 1990). Among
thè crustaceans of thè Kalkschieferzone, besides many
specimens of ostracods, conchostracans (Tintori, 1990)
and of a new species of a “mysidacean” belonging to
Schimperella, Bill 1914 (currently under study) only one
isolated specimen of decapod has been discovered. The
specimen informally named here “ Antrimpos ” sp. n. was
collected from thè middle levels of thè Kalkschieferzone,
in thè quarry of Ca’ del Frate which provided thousands of
conchostracans and Schimperella. This specimen exhibits
features considered by Glaessner (1969) to be diagnostic
of Antrimpos nevertheless, they are quite common in thè
basai penaeids, so their value could be limited. Since thè
erection of Antrimpos by Miinster (1839), many species
of Lower Triassic to Upper Jurassic decapods, often
known by incomplete specimens, were ascribed to this
genus. Also Glaessner (1969), as other authors before him
(e.g. Van Straelen, 1925), considered Antrimpos a collec-
tive genus, to which many Permo-Triassic to Cretaceous
penaeids, not strictly related to modem penaeids, were
assigned. As pointed out by several authors (e.g. Ga-
rassino & Teruzzi, 1993) thè status of this genus should
be revised. Garassino & Teruzzi (1995) assigned thè
Induan (Early Triassic) A. madagascariensis, thè most
ancient species previously attributed to Antrimpos , to
thè new genus Ifasya. More recently Schweigert (2001)
described evolutionary trends in different populations of
Antrimpos from thè Kimmeridgian and Tithonian (Upper
Jurassic) levels of southern Germany, and considered thè
denticulation of thè rostrum and thè development of thè
pereiopods as diagnostic features of thè rank of species.
“ Antrimpos ” sp. n. from thè Kalkschieferzone differs
from Antrimpos noricus Pinna, 1974, from thè Norian
(Upper Triassic) Zorzino Limestone (northem Italy) in
thè larger fourth and fifth pleonites that in A. noricus are
Figs. 1 - A) “ Antrimpos ” sp. n. from thè Meride Limestone. B) Pereiopods.
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
69
very reduced, in thè absence of backward protrusion in thè
posterior margins of pleonites and also in thè shortest first
pleonite. Antrìmpos atavus. Bill, 1914, from thè Anisian
(Middle Triassic) of thè northem Vosges, is only known
in dorsal preservation (Bill, 1914; Gali, 1971); neverthe-
less, “ Antrimpos ” sp. n. differs from it in thè presence of
a gastro-frontal and dorsal ridge, and thè more developed
pereiopods have robust podomeres. The specimen differs
also from Antrimpos mirigiolensis Etter, 1994, from thè
upper Anisian-lower Ladinian Grenzbitumenzone (Monte
S. Giorgio, Switzerland), due to thè omaments of thè
carapace, like thè prominent postocular spine, thè long
spiny rostrum and thè different proportions of pleonites.
The poorly known Antrimpos crassipes (Bronn, 1858)
exhibits chelae with bulbous propodi.
References
Bill P. C., 1914 - Uber Crustaceen aus dem Voltziensand-
stein des Elsasses. Mittheilungen der Geologischen
Landesanstalt Elsass-Lothringen, 8: 289-338.
Briggs D. E. G. & Gali J. C., 1990 - The continuum in
soft-bodied biotas from transitional environments: a
quantitative comparison of Triassic and Carboniferous
Konservat-Lagerstatten. Paleobiology, Lawrence, 16
(2): 204-218.
Gali J. C., 1971 -Faunes etpaysages du Grès à Voltzia du
Nord des Vosges. Essai paléoécologique sur le Bunt-
sandstein supérieur. Mémoire Service Carte géologi-
que Alsace Lorraine, Strasburg, 34: 1-318.
Garassino A. & Teruzzi G., 1993 - A new decapod crus-
tacean assemblage from thè Upper Triassic of Lom-
bardy (N. Italy). Paleontologia Lombarda, Nuova
Serie, Milano, 1: 1-27.
Garassino A. & Teruzzi G., 1995 - Studies on thè Permo-
Trias of Madagascar. 3. The decapod crustaceans of thè
Ambilobè region (NW Madagascar). Atti della Società
italiana di Scienze naturali e del Museo civico di Storia
naturale di Milano, Milano, 134 (1): 85-113.
Glaessner M. F., 1969 - Decapoda. In: Treatise on Inver¬
tebrate Paleontology. Part. (R). Arthropoda 4. R. C.
Moore (ed.). Geological Society of America, Law¬
rence: R399-R651.
Miinster G., 1839 - Decapoda Macrura. Abbildungen und
Beschreibung der fossilen langschwànzeigen Krebse
in den Kalkschiefem von Bayem. Beitràge zur Petre-
factenkunde, Bayreuth, 2: 1-88.
Schweigert G., 2001 - Eine neue Art der Gattung Antrim¬
pos Miinster (Crustacea, Decapoda, Penaeidae) aus
dem Oberjura Siiddeutschlands. Stuttgarter Beitràge
zur Naturkunde, Stuttgart, B, 307: 1-33.
Senn A., 1924 - Beitràge zur Geologie des Alpensiidran-
des zwischen Mendrisio und Varese. Eclogae geologi-
cae Helvetiae, 18: 550-632.
Tintori A., 1990 - Estherids from thè Kalkschieferzone
(Triassic) of Lombardy (N. Italy). Atti del Quarto
Simposio di Ecologia e Paleoecologia delle Comu¬
nità Bentoniche. Sorrento 1-5 Novembre 1988.
Museo Regionale di Scienze Naturali, Torino: 95-
105.
Van Straelen V., 1925 - Contribution a l’étude de Crus-
tacés Décapodes de la période Jurassique. Mémoires
de l’Académie royale Belgique, 7.
Cristiano Larghi - ENI S.p.A., E. & P. Division, Via Emilia 1, 20097 S. Donato Milanese (Milano), Italy.
e-mail: cristiano.larghi@eni.it
Andrea Tintori - Dip. Scienze della Terra “Ardito Desio”, Università degli Studi di Milano, Via Mangiagalli 34, 20133 Milano, Italy.
e-mail: andrea.tintori@unimi.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Sergio Marangon & Antonio De Angeli
Preservation of some specimens of Portunus monspeliensis
(A. Milne Edwards, 1860) from thè Miocene of Sardinia (Italy)
The Cenozoic decapods from Sardinia have been
described by Meneghini (1857), A. Milne Edwards
(1860), Parona (1887), Mariani & Parona (1887), Ristori
(1888), Lovisato (1902), Lòrenthey (1909), Comaschi
Caria (1950, 1956), Marras & Ventura (1991), and De
Angeli & Marangon (1992). The studied specimens come
from marly limestone or calcareous marls from thè Oli¬
gocene, Miocene, and Pliocene.
Portunus monspeliensis (A. Milne Edwards, 1 860) is
usually common in thè middle Miocene levels of Sar¬
dinia and it has been reported by several authors with dif-
ferent synonymies: Lupa bastata Linnaeus, 1767, Nep-
tunus convexus Ristori, 1888, N. granulatus (A. Milne
Edwards, 1860) from Cagliari (Monte S. Michele and
Fangario), Oristano (Pianu, Magomadas, Flussio, and
Tresnuraghes), and Sassari (Bonorva) (Comaschi Caria,
1956). This species is also known from thè Miocene of
Lecce (Puglia), S. Maria Vigliana (Emilia-Romagna),
and Meduno (Friuli-Venezia Giulia) (Ristori, 1888; De
Angeli & Garassino, 2006). The presence of P. monspe¬
liensis from thè Rupelian (lower Oligocene) of Bacino
Ligure Piemontese is dubious. In fact, Allasinaz (1987)
pointed out that thè specimens assigned to this species,
discovered in Piedmont, could belong to a different
subspecies within P. monspeliensis. Finally, this species
was also reported from Malta, Spain, Portugal, France,
Hungary, Austria, Egypt, and Sinai Peninsula (A. Milne
Edwards, 1860; Ristori, 1888; Glaessner, 1928, 1933;
Lòrenthey & Beurlen, 1929; Miiller, 1978, 1984, 1993;
Gatt, 2006).
The descriptions and illustrations of this species
usually have been made based upon carapaces without
exocuticle and therefore thè dorsal surface has not thè
originai omamentation, but a surface with small ovoid
tubercles weakly raised that are present in thè outer part
of thè endocuticle.
The specimens from thè marly limestone of thè
Langhian (middle Miocene) of Binu Mancu (thè area
among thè villages of Flussio-Tresnuraghes-Magoma-
das) provided new contributions to thè knowledge of
this species. In fact, these specimens show thè following
morphological characters: originai dorsal omamenta¬
tion; exocuticle with small granulations, more frequent
on metagastric, cardiac, and branchial regions; elongate
supraorbital margin and interrupted by two narrow fis-
sures and finely granulate (Fig. 1 D); extraorbital spine
more developed than thè other seven spines of lateral
margin; anterolateral spines decrease in sizes (excluding
thè elongate spine located on thè angle); small granula¬
tions also present on thè margins of thè lateral spines;
protogastric regions with transverse granulate crista;
epibranchial regions crossed by a sinuous, granulate
and transverse crista extending also onto thè elongate
spine of thè anterolateral angle; posterolateral margin
concave and with superficial granulate crista; stemite IV
with transverse concavity and finely granulate anterior
lateral margin (Fig. 1 F). The exocuticle of thè carapace,
chelipeds, stemites, and abdomen is always of brown-
purple colour; many small areas white-yellow in colour
are present on chelipeds and surface of carapace (Fig.
1 A-B-C). These areas are larger on thè posterior part
of thè back and smaller and uniform on its other parts.
The carapace is brown-purple in colour and thè areas are
reduced or absent in other specimens. One specimen has
thè carapace with white exocuticle and chelipeds with
brown-purple colour and small areas with white-yellow
colour (Fig. 1 G-H).
The colouring of P. monspeliensis is similar to that of
some Recent portunids, widespread along thè American
coasts (e. g. Arenaeus cribrarius (Lamarck), Arenaeus
mexicanus (Gerstaecker) exhibiting a carapace of grey
or brown colour with small yellow or white areas. The
fossil specimens with white carapace (Fig. 1 G) could be a
case of albinism. In fact, Rathbun (1930, p. 104) reported
occasionai examples of albinism in Callinectes sapidus
Rathbun.
The new specimens of P monspeliensis from thè
Miocene of Sardinia represent an “ unicum ” in thè fossil
record because they have preserved thè chromatic charac¬
ters, useful to observe thè variations of colour within thè
same species.
References
Allasinaz A., 1987 - Brachyura decapoda oligocenici
(Rupeliano) del Bacino Ligure Piemontese. Bollettino
del Museo regionale di Scienze naturali di Torino,
Torino, 5 (2): 509-566.
Comaschi Caria I., 1950 - Crostacei Decapodi nel
Miocene (Elveziano) di Bosa in Sardegna. Ren¬
diconto del Seminario della Facoltà di Scienze
dell’Università di Cagliari, Cagliari, 20 (3-4): 324-
327.
Comaschi Caria I., 1956 - I Crostacei miocenici della
Sardegna. Bollettino del Servizio Geologico d’Italia,
Roma, 78 (1-2): 283-290.
3™ SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
71
Fig 1 - A-B-C) Portunus monspeliensis A. Milne Edwards, 1860, n. cat. MSNM Ì26874. A-B) Dorsal view. C) Cheliped. D-E)
P. monspeliensis A. Milne Edwards, 1860, n. cat. MSNM Ì26875. D) Anterior margin. E) Dorsal view. F) P. monspeliensis A.
Milne Edwards, 1860, n. cat. MSNM Ì26876, ventral view. G-H) P. monspeliensis A. Milne Edwards, 1860, n. cat. MSNM
Ì26877. G) Dorsal view. H) Cheliped.
72
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
De Angeli A. & Marangon S., 1992 - Necronectes schaf-
feri Glaessner nel Miocene della Sardegna. Lavori
- Società Veneziana di Scienze Naturali , Venezia, 17:
175-182.
De Angeli A. & Garassino A., 2006 - New reports of
decapod crustaceans from thè Mesozoic and Cenozoic
of Friuli-Venezia Giulia (NE Italy). Atti della Società
italiana di Scienze naturali e del Museo civico di
Storia naturale in Milano , Milano, 147 (2): 267-294.
Gatt M., 2006 - Il-Geologija u 1-Paleontologija tal-Gzejjer
Maltin - 1. Pubblikazzjonijiet Indipendenza , Malta.
Glaessner M. F., 1928 - Die Dekapodenfauna des òster-
reichischen Jungtertiars. Jahrbuch Geologie Bunde-
sanstein, Wien, 78 (2): 161-219.
Glaessner M. F., 1933 - New Tertiary Crabs in thè Collec-
tion of thè British Museum. TheAnnals and Magazine
of Naturai History, London, voi. XII, Tenth series, 67:
1-28.
Lòrenthey I. E., 1909 - Beitràge zur tertiàren Dekapo¬
denfauna Sardiniens. Mathematischen und Natunvis-
senschaftlichen Berichte aus Ungarn, Budapest, 24:
202-257.
Lòrenthey I. E. & Beurlen K., 1929 - Die Fossilen Deka-
poden der Lànder der ungharischen Krone. Geologica
Hungarica, Serie Paleontologica, Budapest, 3: 1-420.
Lovisato D., 1902 - Le specie fossili finora trovate nel
calcare compatto di Bonaria e S. Bartolomeo. Tipo-
Litografia Commerciale, Cagliari, 1-21.
Mariani E. & Parona C. F., 1887 - Fossili Tortoniani di
Capo S. Marco in Sardegna. Atti della Società italiana
di Scienze naturali, Milano, 30: 101-191.
Marras G. & Ventura G., 1991 - Crostacei decapodi del
Miocene di Sassari (Sardegna nord-occidentale). Bol¬
lettino della Società Sarda di Scienze Naturali, Sas¬
sari, 28: 105-119.
Meneghini G.,1857 - Paléontologie de File de Sar-
daigne. In: Voyage en Sardaigne, Ed. La Marmora,
Torino.
Milne Edwards A., 1860 - Monographie des Décapodes
macrures fossiles de la famille des Thalassiniens.
Annales des Sciences Naturelle Zoologie, Paris, sér.
4, 14: 294-257.
Miiller P., 1978 - Decapoda (Crustacea) fauna a budapesti
miocénbòl (5). Fòldtani Kòzlòny, Budapest, 108: 272-
312.
Miiller P., 1984 - A badeni emelet tizlàbù ràkjai. Decapod
Crustacea of thè Badenian. Geologica Hungarica,
Serie Paleontologica, Budapest, 42 : 1-121.
Miiller P., 1993 - Neogene Decapod Crustaceans from
Catatonia. Scripta Musei Geologici Seminarii Barci-
nonensis, Barcelona, 225: 1-39.
Parona C. F., 1887 - Appunti per la Paleontologia mio¬
cenica della Sardegna. Bollettino Società Geologica
Italiana, Roma, 6: 287-358.
Rathbun M. J., 1930 - The cancroid crabs of America
of thè families Eurylidae, Portunidae, Atelecyclidae,
Cancridae and Xanthidae. United States National
Museum Bulletin, Washington, 152: 1-593
Ristori G., 1888 - Alcuni Crostacei del Miocene medio
italiano. Atti della Società Toscana di Scienze Natu¬
rali, Pisa, 9: 212-219.
Sergio Marangon - Via Anemone 8, 20090 Segrate (Milano), Italy.
e-mail: Sergiomarangon@libero.it
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Sergio Marangon & Antonio De Angeli
New decapod assemblage from thè lower Oligocene (Rupelian)
of Bacino Ligure Piemontese (NW Italy)
The fossil crustaceans from Bacino Ligure Piemon¬
tese have been described by several authors. E. Sismonda
(1846, 1861) reported some decapods from thè Miocene
and Pliocene of Asti and Torino. Later, Michelotti (1861),
Crema (1895), and above all Ristori (1886, 1889), stud-
ied extensively thè Oligocene brachyurans of this region.
Recently, Allasinaz (1987), Marangon & De Angeli
(1997), De Angeli & Marangon (2001, 2003), and Larghi
(2003) provided new contributions to thè knowledge of
thè decapod fauna from thè Oligocene.
The Oligocene rocks of Bacino Ligure Piemontese lie
on a metamorphic formation of Continental origin without
fossils (Brecce di Cravara). Conglomerates with fossils
of plants (Formazione di Pianfolco) are intercalated with
these rocks. Transgressive deposits rich in fossils from
thè middle Rupelian (Formazione di Molare) lie on these
conglomerates (Charrier, Femandez & Malaroda, 1964;
Franceschetti, 1967; Bianco, 1985; Balossino & Bianco,
1986; Fantoni, Lovati & Rossi, 1983).
The new specimens (Marangon & De Angeli, work
in progress) preserved in nodules of diagenetic origin,
come from thè middle Rupelian levels (Formazione di
Molare) of Case Cherpione (Alessandria). The studied
specimens are referred to Lophoranina aculeata (A.
Milne Edwards, 1881) (Raninidae De Haan, 1839),
Ethusa n. sp. (Dorippidae MacFeay, 1838), Retropluma
n. sp. (Retroplumidae Gill, 1894), Palaeocarpilius aqui-
tanicus A. Milne Edwards, 1862 (Carpiliidae Ortmann,
1893), and Asthenognathus n. sp. (Pinnotheridae De
Haan, 1833).
The presence of Lophoranina from thè Oligocene of
Bacino Ligure Piemontese is already known by two cara-
paces that E. Sismonda (1861) ascribed to Lophoranina
aldrovandii (Ronzani, 1818). The comparison with thè
holotype of L. aldrovandii , housed in thè Museo di Pa¬
lazzo Poggi, Bologna, and thè holotype of L. aculeata (A.
Milne Edwards, 1881), housed in thè Museum national
d’Histoire naturelle in Paris, permitted assigning one
new specimen and those described by E. Sismonda to L.
aculeata.
Ethusa n. sp. (Marangon & De Angeli, work in
progress) has morphological affinities with E. chibai
Karasawa, 1993, from thè lower Pliocene of Japan which
has a narrower cardiac region and less deep branchial
grooves. Miiller (1984) and Via Boada (1988) reported
thè presence of one probable new subspecies of Ethusa
mascarone (Herbst, 1785) from thè Messinian (Miocene)
of Santa Pola (Alicante, Spain). Ethusa n. sp. represents
1 thè oldest species known to date in this genus.
Retropluma Gill, 1894, is known from three fossil
species: R. eocenica Via Boada, 1959 (Ilerdian and Lute-
tian - Spain and Italy); Retropluma laurentae Collins,
Lee, Noad, 2003 (Miocene and Pleistocene - Sabah and
Sarawak); R. craveri (Crema, 1895) (Pliocene - Italy).
Retropluma n. sp. (Marangon & De Angeli, work in
progress) differs from thè other species in thè shape of
thè carapace and thè ornamentation of thè dorsal cari-
nae.
The assignment to Palaeocarpilius macrochelus
(Desmarest, 1822) of thè specimens described by Ris¬
tori (1889) and Allasinaz (1987) is dubious. The recent
review of thè fossil species of Palaeocarpilius by
Beschin & De Angeli (2006) pointed out thè morpho¬
logical characters that distinguish thè Eocene specimens
of P macrochelus (Desmarest, 1822) and thè Oligocene
specimens of P. aquitanicus A. Milne Edwards, 1862,
by a comparison among thè holotypes, housed in thè
Museum national d’Histoire naturelle, Paris. The new
specimen of Case Cherpione is ascribed to P. aquitani¬
cus because it exhibits seven anterolateral lobes instead
of eight as in thè specimens of P. macrocheilus (Desmar¬
est, 1822).
Asthenognathus Stimpson, 1858, is known from three
Recent and five fossil species: A. cornishorum Schweitzer
& Feldmann, 1999 (lower Miocene - United States); A.
globosa (Karasawa, 1990) (lower Miocene - Japan); A.
microspinus Casadio, De Angeli, Feldmann, Garassino,
Hetler, Parrai & Schweitzer, 2004 (Oligocene - Argen¬
tina); A. urretae Schweitzer & Feldmann, 2001 (upper
Oligocene - Argentina); A. laverdensis De Angeli & Ga¬
rassino, 2006 (lower Oligocene - Italy). Asthenognathus
n. sp. (Marangon & De Angeli, work in progress) has a
smooth and subrectangular carapace, differing from A.
laverdensis in having almost parallel lateral margins and
more prominent anterolateral angles (De Angeli & Ga¬
rassino, 2006).
The faunistic assemblage of Bacino Ligure Piemon¬
tese is strictly correlated with thè Rupelian faunae of
Vicenza (Italy) and Biarritz (France). To date, thè spe¬
cies described are thè following: Hoploparia sp.; Cal-
lianassa sp.; Pagurus sp.; Callianassa canavari Ristori,
1889; Zygopa galantensis (De Angeli & Marangon,
2001); Ranina speciosa (Munster, 1840); Lophoranina
aculeata (A. Milne Edwards, 1881); Ethusa n. sp.
(Marangon & De Angeli, work in progress); Calappa
sp.; Calappilia mainii Allasinaz, 1987; C. vicentina
Fabiani, 1910; C. verrucosa A. Milne Edwards, 1873;
Mursiopsis postulosus Ristori, 1889; Cherpiocarcinus
rostratus Marangon & De Angeli, 1997; Retropluma n.
sp. (Marangon & De Angeli, work in progress); Palaeo¬
carpilius aquitanicus A. Milne Edwards, 1860; Portu-
nus convexus (Ristori, 1889); Coeloma vigil A. Milne
Edwards, 1865; Asthenognathus n. sp. (Marangon & De
Angeli, work in progress).
j
«
74
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Fig. 1 - A) Lophoranina aculeata (A. Milne Edwards, 1881), n. cat. MSNM Ì26879. B) Ethusa n. sp., n. cat. MSNM Ì26880.
C) Retropluma n. sp., n. cat. MSNM Ì26881. D) Asthenognathus s. nov., n. cat. MSNM Ì26882. E) Palaeocarpilius aquitanicus A.
Milne Edwards, 1860, n. cat. MSNM Ì26883.
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
75
References
Allasinaz A., 1987 - Brachyura Decapoda oligocenici
(Rupeliano) del Bacino Ligure Piemontese. Bollettino
del Museo regionale di Scienze naturali di Torino,
Torino, 5 (2): 509-566.
Balossino P. & Bianco P., 1986 - Nota preliminare sulla
biostratigrafia dell’area di Ponzone, Cimaferle e Ban¬
dita (Piemonte SE). Bollettino del Museo regionale di
Scienze naturali di Torino, Torino, 4 (2): 469-481.
Beschin C. & De Angeli A., 2006 - Il genere Palaeo-
carpilius A. Milne Edwards, 1862 (Decapoda, Bra¬
chyura, Carpiliidae) nel Terziario del Vicentino (Italia
settentrionale). Studi e Ricerche - Associazione Amici
del Museo - Museo Civico “G. Zannato ”, Montecchio
Maggiore (Vicenza), 13: 5-17.
Bianco P., 1985 - Nota preliminare sulla biostratigrafia
dell’area di Spigo Monferrato, Pareto e Mioglia (Alpi
Liguri, Italia N.O.). Atti Accademia Nazionale dei
Lincei, Roma, 8, 78: 34-43.
Charrier G., Femandez D. & Malaroda R., 1964 - La
formazione di Pianfolco (Bacino oligocenico Ligure
Piemontese). Atti Accademia Nazionale dei Lincei,
Roma, 87 (2): 25-82.
Crema C., 1895 - Sopra alcuni decapodi terziari del
Piemonte. Atti della Reale Accademia di Scienze di
Torino, Torino, 30: 664-681.
De Angeli A. & Garassino A., 2006 - Asthenognathus
laverdensis n. sp. (Decapoda: Brachyura: Pinnotheri-
dae) firom thè lower Oliogecene of Laverda (Vicenza,
NE Italy). Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale in Milano,
Milano, 147 (2): 295-304.
De Angeli A. & Marangon S., 2001 - Paralbunea
galantensis, nuova specie di anomuro oligocenico
del Bacino Ligure-Piemontese (Italia settentrionale).
Studi Trentini di Scienze Naturali, Acta Geologica,
Trento, (1999), 76: 99-105.
De Angeli A. & Marangon S., 2003 - Contributo alla con¬
oscenza dei Decapodi oligocenici del Bacino Ligure
Piemontese (Italia settentrionale). Atti della Società
italiana di Scienze naturali e del Museo civico di
Storia naturale in Milano, Milano, 144 (2): 185-196.
Fantoni R., Lovati I. & Rossi P. M., 1983 - La successione
oligocenica tra Ovada e Cassinelle (AL). Evoluzione
paleogeografia ed implicazioni strutturali. Rivista Ital¬
iana di Paleontologia, Roma, 88 (2): 251-270.
Franceschetti B., 1967 - Studi geologici sulla regione ad
Ovest di Ovada (Provincia di Alessandria). Memorie
della Società Geologica Italiana, Roma, 6: 379-420.
Larghi C., 2003 - First record of Oligocene retroplumid
crab (Crustacea: Decapoda: Brachyura) from Italy.
Bulletin of thè Mizunami Fossil Museum, Mizunami,
30: 57-60.
Marangon S. & De Angeli A., 1997 - Cherpiocarcinus a
nuovo genere di brachiuro (Decapoda) dell’Oligocene
del Bacino Ligure-Piemontese (Italia settentrionale).
Lavori - Società Veneziana di Scienze Naturali, Ve¬
nezia, 22: 97-106.
Michelotti G., 1861 - Etudes sur le Miocène infèrieur de
l’Italie septentrionale. Memoire de la Société Hollan-
dais des Sciences, Harlem.
Mtiller P., 1984 - Messinian and older decapods from
Mediterranean with description of two new species.
Ann. Geol. Pays Helleniques, Athens, 32: 25-34.
Ristori G., 1886 - I Crostacei Brachiuri e Anomuri del
Pliocene italiano. Bollettino della Società Geologica
Italiana, Roma, 5: 93-129.
Ristori G., 1889 - I Crostacei Piemontesi del Miocene
inferiore. Bollettino della Società Geologica Italiana,
Roma, 7: 397-413.
Sismonda E., 1846 - Descrizione dei Pesci e Crostacei
fossili nel Piemonte. Memorie della Reale Accademia
di Scienze di Torino, Torino, 2(10): 1-89.
Sismonda E., 1861 - Appendice alla descrizione dei Pesci
e Crostacei fossili del Piemonte. Memorie della Reale
Accademia di Scienze di Torino, Torino, 2(19): 1-24.
Via Boada L., 1988 - Els decàpodes. In: Història Naturai
dels Pai'sos Catalans, Enciclopédia Catalana, S.A.,
Barcellona, 15: 343-352.
Sergio Marangon - Via Anemone 8, 20090 Segrate (Milano), Italy.
e-mail: Sergiomarangon@libero.it
Antonio De Angeli - Associazione Amici del Museo “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy.
e-mail: antonio_deangeli@virgilio.it
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
i
Rafael Gioia Martins-Neto & Sebastiào Carlos Dias Iùnior
The Brazilian paleodecapod fauna: state of thè knowledge
The decapod fauna is well represented in Brazilian
deposits with several described species, especially from
Cenozoic sediments. Mesozoic decapods are known
from thè Riachuelo Formation (Lower Cretaceous, Ser-
gipe), Santana Formation (Lower Cretaceous, Cearà),
and Itamaracà and Gramame formations (Lower and
Upper Cretaceous respectively, Pemambuco). Cenozoic
decapods are known from thè Maria Farinha Formation
(Paleocene, Pemambuco), Cicero Dantas Formation
(Eocene/Oligocene, Bahia), Pirabas Formation (Miocene)
and Tremembé Formation (Oligocene, Sào Paulo) (see
Martins-Neto, 2005). Recently new species were added:
in thè Riachuelo Formation (Albian, Sergipe basin) frag-
ments attributable to palinurans were recorded (Reis et
al., 2005). Reis et al. (2005) suggested a dose affìnity of
one specimen to thè family Scyllaridae, and they note as
thè diagnostic characteristic thè presence of three carinae
on thè cephalic region. Just two families, Coleiidae and
Polychelidae, within thè Eryonoidea, exhibit three par-
allel carinae on thè dorsal carapace surface, however,
neither have recorded genera for thè Cretaceous. So, until
more complete material is known, this specimen must
be considered Eryonoidea family, genus, and species
undetermined, possibly new. Apart from this, two spe¬
cies attributed to thè genus Dardanus were reported in
thè Paleocene sediments of thè Maria Farinha Formation
(Tàvora et al., 2005). For thè Pirabas Formation Calap-
pilia brooksi Ross & Scolaro, and thè species Arenaeus
cribarìus (Lamarck) and Glyphithyreus sturgeoni Feld-
mann et al. (Tàvora et al., 2005) was recorded. A com¬
plete list of thè known Brazilian decapods is fumished in
Table I, updated from Martins-Neto (2005).
Table 1 - Summary of thè Brazilian paleocarcinofauna (updated from Martins-Neto, 2005).
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
77
The Santana Formation crustaceans (Cretaceous,
northeast Brazil) are preserved in an excellent state
of preservation, several are complete, articulated, and
tridimensional. The fossilization process is due to iron
oxide substitution (oxidation) and more rarely apa-
;; tite (phosphatization). The environment in this case is
! lacustrine. Another type of lacustrine environment as in
t thè Tremembé Formation (Oligocene, southeast Brazil)
isolated cephalothorax pieces as well as fragments
of abdomen are found, indicative of several stages of
fragmentation. In some specimens thè pereiopods stili
remain articulated, but most parts of thè specimens are
compacted in thè sediment (pirobituminous shale) and thè
main fossilization process is of thè quitine substitution
and calcium carbonates by pyrite or marcasite (pyritiza-
tion). In environments with some marine influence, as is
thè case of thè Romualdo Member (concretion level of thè
Santana Formation), thè crustaceans are fragmented (iso¬
lated pieces of cephalothorax and rare abdomen pieces).
They are totally disarticulated and preserved as internai
moulds. In this specific case, thè individuai accumula-
tion in a concretion is suggestive of an environmental
stress. In thè Pirabas Formation (Miocene, north Brazil),
thè environment is marine and isolated carapaces, some
pereiopod, fragments, brachyurans chelipeds, isolated
carapaces of lobster-like crustaceans and well preserved
pieces in concretions are found. The fossilization process
is variable, ffom moulds to partially permineralized speci¬
mens. The majority of these specimens are tridimensional,
not articulated and in final steps of fragmentation.
78
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
References
Martins-Neto R. G., 2005 - Estàgio atual da paleoartropo-
dologia brasileira: hexàpodes, miriàpodes, crustàceos
(Isopoda, Decapoda, Eucrustacea e Copepoda) e
quelicerados. Arquivos do Museu Nacional, Rio de
Janeiro, 63 (3): 471-494.
Reis M. A. F., Turbay C. V. E. & De Ceserò P., 2005 -
Deserto de um novo Decapoda (Natantia, Malaco-
straca, Crustàcea) da Formando Riachuelo, Albiano da
Bacia de Sergipe. Armàrio do Instituto de Geociéncias
- UFRJ, 28 (1): 80-91.
Tàvora V. A., Miranda V. F. O., Viegas L. G. F. & Galvào
P. H. F., 2005 -Novos registros de crustàceos decàpo-
des do Cenozóico (Paleoceno e Mioceno) do Brasil.
Revista Brasileira de Geociéncias , 35 (3): 393-400.
Rafael Gioia Martins-Neto - Universidade Federai de Juiz de Fora, Campus Universitario, Martelos, 36036-900-Juiz de Fora, MG Brazil.
e-mail: martinsneto@terra.com.br
Sebastiào Carlos Dias Junior - Universidade Federai de Juiz de Fora, Campus Universitario, Martelos, 36036-900-Juiz de Fora, MG Brazil.
e-mail: scdjunior@click21.com.br
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Paolo Monaco, Jesùs E. Caracuel, Alice Giannetti, Jesùs M. Soria &
Alfonso Yébenes
Thalassinoides and Ophiomorpha as cross-facies trace fossils
of crustaceans from shallow-to-deep-water environments:
Mesozoic and Tertiary examples from Italy and Spain
The morphological expression of trace fossils varies
according to whether they are preserved, within/between
sandstone or mudstone beds. For this reason a toponomy
has been proposed and a series of terms, thè so-called
Seilacher’s and Martinsson’s terms, has arisen to describe
thè preservation potential of trace fossils. They are:
hyporelief or hypichnia (base of bed at boundary), full
relief or endichnia (inside bed), full relief or exichnia
(outside bed) and epirelief or epichnia (top boundary)
(Seilacher, 1964; Martinsson, 1970). Both of these clas-
sifications involve structures which are strictly related
to thè casting medium of an event bed or transition with
underlying or overlying beds (Bromley, 1990). Typi-
cal expression of hyporelief are graphoglyptids (Fuchs,
1895), a large supergroup of geometrical trace fossils
usually preserved at soles of deep-water turbidite deposits
of basin plains, which includes a very high diversity of
ichnogenera and species (Seilacher, 1977).
Thalassinoides-type branched fossil burrow Systems
in thè geological record, known mainly as Thalassinoides
and Ophiomorpha (chiefly but not exclusively produced
by decapod crustaceans), usually are preserved in beds as
hypichnia, endichnia and exichnia but their preservation
potential and toponomy strictly depend upon thè environ-
ment and type of event sedimentation. Moreover, thè large
distribution of these ichnogenera differ from graphoglyp¬
tids which are strictly confined to deep water conditions
and affecting inter-turbidite muddy deposits (Monaco, in
press). Thalassinoides and Ophiomorpha, therefore, are
typical cross-facies trace fossils (Frey et al., 1978), which
usually spread in every environments during thè Phan-
erozoic and occur from shallow facies (beach, near-shore
and shelf), mainly in thè Paleozoic and Mesozoic, to very
deep environments (slope to distai basin plains) in thè
Tertiary (Ksiqzkiewicz, 1970; Frey et al, 1978; Palmer,
1978; Archer & Maples, 1984; Sheehan & Schiefelbein,
1984; Bromley, 1990; Fiirsich, 1998; Uchman, 1998;
Monaco, 2000b; Monaco & Garassino, 2001; Tchou-
matchenco & Uchman, 2001; Giannetti & Monaco,
2004). The ichnogenus Thalassinoides Ehrenberg (1844)
is applied to smooth-walled, essentially cylindrical Y- to
T-shaped branched trace fossils. In thè Early Jurassic
time Thalassinoides suevicus (Rieth 1932) characterizes
shallow-marine environments as three-dimensional Sys¬
tems of branched, Y-shaped tunnels of variable diameter
enlarged at thè bifurcation points (Fiirsich & Oschmann,
1993; Giannetti & Monaco, 2004). This form is pre-
dominantly horizontal, more-or-less regularly branched,
essentially cylindrical, and dichotomous bifurcations are
more common than T-shaped branches (Howard & Frey,
1984). The origin and palaeoenvironmental significance
of Thalassinoides has been recently summarised (Howard
& Frey, 1984; Ekdale, 1992). Instead, thè ichnogenus
Ophiomorpha Lundgren (1891) commonly occurs as
vertical to horizontal or sub-horizontal cylindrical tunnel
systems, seldom branched and covered by elongate or
irregular pellets arranged perpendicular to thè long axis.
Branching points are usually preserved in both ichnotaxa
but these features are more typical of some ichnospecies
(e.g. T. suevicus) and reveal sharp angles, locally with
characteristic enlargements (Monaco & Giannetti, 2002;
Giannetti & Monaco, 2004). Some horizontal segments
of Ophiomorpha, lacking knobby appearance, resemble
typical Thalassinoides, but vertical portions are generally
lined (Frey et al., 1978; Uchman, 1995; Monaco, 2000).
These trace fossils have been generally attributed to thè
activity of crustaceans, but not only decapods (Frey et al.,
1978; Monaco & Garassino, 2001), and their toponomic
preservation is here investigated in four cases of event
sedimentation from Mesozoic and Cenozoic deposits.
The four case-studied are thè follow: A) Lower
Jurassic (Pliensbachian) lagoonal limestones and marly
beds of thè northeastem Prealps (Calcari Grigi); B)
Lower Cretaceous inner shelf with unidirectional trac-
tive contour currents, from eastem Spain (Serra Gelada);
C) Lower Jurassic outer shelf to basinal nodular deposits
with storm/turbidite calcarenites, centrai Italy (Rosso
Ammonitico); D) Miocene slope to basin plain systems
with deep water turbidite deposits, northem Apennines
(Macigno and Marnoso Arenacea flysch deposits).
A) In lagoonal nodular limestones of thè Early Jurassic
(Pliensbachian) in northeastem Prealps (Calcari Grigi
formation), trace fossils of crustaceans ( Thalassinoides
and Ophiomorpha) are very abundant. They are found in
limestones forming hexagonal mazes distributed horizon-
tally, and they are particularly well developed at thè base
of thè recorded parasequences: endichnia and hypichnia.
In thè well-exposed endichnia and hypichnia thè maze is
regular in shape, branching points exhibit enlargements
and thè diameter of tunnels decreases progressively from
thè branched points (diameter varies from 16 to 4 cm).
Meshes are horizontal, forming irregular cells up to 30
cm wide, and locally tunnels are randomly distributed in
thè marls and at thè limestone/marl transition at thè base
of thè parasequences. In this case, bioturbation can reach
down thè marly levels producing nodules suggesting that
thè trace maker was active to maximize thè capture of
organic material when thè substratum has a low nutri-
L
80
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
tional value (see Giannetti et al, 2007). When tunnels
are in thè clay or marly levels they are particularly flat-
tened (rare case). Crustacean trace fossils develop at thè
base of parasequences during rising sea level, indicating
that oxygenated and nutrient-rich open marine condi-
tions were induced progressively and are well exploited
by crustaceans. In these open marine conditions thè high
rate of sedimentation, oxygenation, and nutrients are ideal
conditions for crustacean burrowing. This new opportun-
istic fauna populates thè lagoon, replacing thè oligo-
trophic fauna (only infaunal bivalves) of restricted and
poorly oxygenated conditions typical of thè low sea level
stand. The arrangement of skeletal grains and peloids in
wall structures of thalassinoideans commonly reflects an
ichnofabrics of decapod crustacean activity (Giannetti
& Monaco, 2004). Structure of thè burrow linings and
lithifìed calcareous walls (see type IV) display concentric
layering due to crustacean and biogenic stria (see type
III and type B of authors), probably related to activity of
decapods (Frey et al., 1978; Hasiotis & Mitchell, 1993;
Anderson & Droser, 1998; Monaco & Garassino, 2001).
In some cases thè shape of some non-branched burrows
( Thalassinoides ? isp.) resembles those of modem sto-
matopods (Monaco & Giannetti, 2002).
B) In thè Lower Cretaceous (Albian) of thè Sàcaras
Formation in thè Serra Gelada succession (Prebetic of
Alicante, Spain) carbonate-rich, upward thickening inner
shelf parasequences were deposited. They are domi-
nated by deposits and features indicating strong currents
(maybe contour currents), and many types of trace fossils
have been identified (Monaco et al, 2005). Shell-cov-
ered, structured trace fossils ( Ereipichnus geladensis )
and unstructured, non-imbricated ones such as Scolicia
and Cardioichnus (produced probably by irregular or
heart-shaped spatangoid sea-urchins) and various types
of Thalassinoides and Ophiomorpha have been recorded
(Monaco et al., 2005). In this section Thalassinoides
consists of branched, Y-shaped hypichnia or endichnia,
similar to those recovered in Calcari Grigi (mainly type II,
III, and IV giant type up to 30 cm in diameter, and type D
similar to Thalassinoides ?isp., see Giannetti et al, 2007).
In some levels a dose relationship exists between shell-
armoured Ereipichnus and large Thalassinoides types.
Horizontal mazes of this branched endichnia are always
distributed deeper (often reaching thè hypichnia position)
than Ereipichnus which tend to occupy shallower tiers
to better utilize strong unidirectional currents at thè sea-
bottom (Monaco et al., 2005). Thalassinoides, therefore,
develops in deeper tiers when currents at sea-floor are
active, and these traces were produced by crustaceans
mainly when high sea level conditions occurred (as in
thè case A). The vertical or oblique Ophiomorpha, con¬
versely, develop as endichnia (more properly crossichnia,
a new category here introduced to describe trace fossils
which cross obliquely two or three beds) exclusively in
calcarenites during thè lowstand intervals at thè top of
parasequences. The Serra Gelada section, therefore, is
important from thè ichnological point of view because
a very dose relationship exists among current sensitive
trace fossils, active bottom currents and sea-level varia-
tions in a shallow water environment.
C) The Toarcian Rosso Ammonitico facies in thè cen¬
trai Apennines (Italy), was deposited in a relatively shal¬
low (100-150m deep) middle-outer shelf to inner basin,
periodically submitted to seaward-oriented tempestites/
turbidites (with hummocky cross-stratification). This
facies is very rich in small and medium size Thalassi¬
noides of types I and II (Monaco et al., 1994; Caracuel
et al., 2000). The medium-size Type II has been found
as hypichnia at thè soles of distai tempestite/turbidite
events and develops when calcarenitic material eroded
and then deposited on thè autochthonous mud (casting).
The small Type I, conversely, is more commonly found
as endichnia (but also epichnia) mainly dose thè top of
a calcarenite beds (calcilutite) and was subjected to tier-
ing, which represents a vertical partitioning of burrow
distribution within thè calcarenitic sediment (Caracuel
et al, 2000). The shallowest, more oxygenated tier of
a typical Rosso Ammonitico tempestile bed, thè nodular
calcilutite interval, is dominated by small Thalassinoides
type I, while other minute trace fossils (e.g. Planolites,
Helminthopsis and Chondrites) are distributed at depth in
thè less oxygenated calcarenite material (Caracuel et al.,
2000). A colonisation depth of tiers therefore reflects thè
new substrate characteristics induced by thè event sedi¬
mentation. Tiering from reduced (base of bed) to more
oxygenated conditions (top of bed) was controlled by thè
substrate characteristics and by thè organic content. The
organic matter may be superfìcially exploited by small
cmstaceans (one ammonite mould exhibits Thalassi¬
noides type I burrow, F. Venturi pers. com.) only when
sedimentation rate of unidirectional/oscillatory flow
regime decreases to normal conditions. When normal
condition re-established (e.g. re-oxygenation) in non-
resedimented marls of Rosso Ammonitico facies, then
small Thalassinoides type I are leaders among other
trace fossils and burrowing of shallower tiers (endichnia/
epichnia) was very abundant inducing nodularity in thè
substrate and producing a “gradational” tiering in marls
(Caracuel et al., 2000).
D) In Oligo-Miocene siliciclastic deep-water turbidite
units of thè northem Appennines (centrai Italy), known as
Macigno, Cervarola and Marnoso Arenacea flysch depos¬
its, trace fossils are very abundant and are represented by
hypichnia, endichnia, exichnia, epichnia, and crossichnia
(if they cross obliquely two or three beds). In thin-bedded,
low-density turbidites graphoglyptids ( Paleodictyon and
many others, see Monaco, in press) are exclusively hyp¬
ichnia, while epichnia groups are formed by totally differ-
ent traces (e.g. Scolicia prisca). Conversely, in medium
to thick-bedded ones (mainly high-density turbidity flow
deposits) Thalassinoides and Ophiomorpha occur as
hypichnia, endichnia exichnia (never epichnia), and cros¬
sichnia when they cross obliquely thè massive sand reach¬
ing thè underlying marly beds. As indicated by Uchman
(1998), thè Ophiomorpha found in flysch deposits differ
from those recovered from shallow water deposits because
they commonly occur as small hypichnial specimens
which are smooth and straight at thè bottom of turbidites
(e.g. O. annidata and O. rectus, see Ksiqzkiewicz (1977).
Large Ophiomorpha rudis (Tunis & Uchman, 2003), is
most common in thè medium to thick bedded turbidites
of Marnoso Arenacea flysch. This very large trace (up to
12 cm in diameter) appears as subhorizontal hypichnia-
endichnia in thè lower half of thè turbidite beds and is
characterized by irregular thick ridges, up to 3-4 cm thick,
commonly developed as large knobby bulges, with irreg¬
ular thickening of tunnel diameter. Locally, branches are
3RD SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
81
very short and represent dead ends. Other forms of high-
density turbidites are unbranched and straight, exhibit-
ing a regular thickening of thè burrow diameters (twice
normal diameter) and producing a typical shape like a car
silencer (car silencer-shaped trace fossils). These forms
are distributed very commonly (30 specimens preserved
at thè BIOSEDLAB of Perugia University) as endich-
nia dose to thè bottom of meter-thick, sandy turbidites,
where they cut giant groove casts at angles up to 60-90°,
suggesting adaptative burrowing strategies to high energy
and tractive current flows. Other Ophiomorpha-Wkc. trace
fossils are preserved as vertical or oblique, cylindrical,
knobby structures, cutting many calcarenite and calcis-
iltite beds (crossichnia). They are unwalled, sand- or silt-
filled, 18-33 mm in diameter reaching up 30 cm in length,
and some exichnia/crossichnia specimens show typical
ring-shaped, transverse segments (Uchman & Demircan,
1999). Others crossichnia are subquadrate in cross sec-
tion, revealing regular knobby surfaces, up to 55 mm in
diameter and 50 cm in length. These structures were prob-
ably not produced by crustaceans. According to (Fòllmi &
Grimm, 1990), thè crustaceans producing Thalassinoides
and Ophiomorpha in deep turbiditic environments may
survive transport by turbiditic currents and can produce
burrows under anoxic conditions for a limited numbers
of days.
The four examples here reported are indicative of
cross-facies thalassinoidean trace fossils which are pro¬
duced by opportunistic organisms. A dose relationship
probably exists among infaunal activity of crustaceans,
event sedimentation, rising of sea level, increasing in thè
oxygen content and characteristics of thè substrate from
shallow to deep water conditions where high-density flow
or peculiar sedimentary processes occur. Further quanti¬
tative analyses are welcome in order to try to define all
relationships between these traces and different cases of
event sedimentation. The fundamental aim is not only to
characterize thè toponomy of these important traces, but
also to define thè palaeoenvironmental features, morpho-
logic expressions (Giannetti et al. , 2007) and distribution
of crustacean activity in marine substrates.
References
Anderson B. G. & Droser M. L., 1998 - Ichnofabrics and
geometrie configurations of Ophiomorpha within a
sequence stratigraphic framework: an example from
thè Upper Cretaceous US western interior. Sedimen-
tology, 45: 379-396.
i Archer A. W. & Maples C. G., 1984 - Trace fossil distri¬
bution across a marine to non marine gradient in thè
Pennsylvanian of South Western Indiana. Journal of
Paleontology, Lawrence, 58: 448-466.
Bromley R. G. C., 1990 - Trace fossils, biology and
taphonomy. Special topics in paleontology. Unwin
Hyman Ltd, London.
Caracuel J., Monaco P. & Olóriz F., 2000 - Taphonomic
tools to evaluate sedimentation rates and stratigraphic
completeness in Rosso Ammonitico facies (epioceanic
tethyan Jurassic). Rivista Italiana di Paleontologia e
Stratigrafia, Milano, 106 (3): 353-368.
Ehremberg K., 1944 - Ergànzende Bemerkungen zu
den seinerzeit aus dem Miozàan von Burgschlein-
itz beschrieben Gangkemen und Bauten dekapoder
Krebse. Palàontologische Zeitschrift, Berlin, 23:
345-359.
ì Ekdale A. A., 1992 - Mudckraking and mudslinging:
thè joys of deposit-feeding. In: Trace fossils. Short
Courses in Paleontology. C. G. Maples & R. R. West
(eds.). The Paleontological Society, Knoxville, Ten¬
nessee.
Fòllmi K. B. & Grimm K. A., 1990 - Doomed pioneers:
Gravity-flow deposition and bioturbation in marine
oxygen-deficient environments. Geology, 18: 1069-
1072.
I Frey R. W., Howard J. D. & Pryor W. A., 1978 - Ophio¬
morpha-. its morphologic, taxonomic, and environmen-
tal signifìcance. Palaeogeography Palaeoclimatology
Palaeoecology, Amsterdam, 23: 199-229.
| Fuchs T., 1895 - Studien iiber Fucoiden und Hiero-
glyphen. Denkshriften der Kaiserlichen Akademie
der Wissenshaften, Wien, Matematisch Naturwissen-
schaftliche, Klasse, 62: 369-448.
Fiirsich F. T., 1998 - Environmental distribution of trace
fossils in thè Jurassic of Kachchh (Western India).
Facies, 39: 243-272.
Fiirsich F. T. & Oschmann W., 1993 - Shell beds as tools
in basin analysis: thè Jurassic of Kachchh, western
India. Journal of thè Geological Society, London,
150: 169-185.
Giannetti A. & Monaco P., 2004 - Burrow decreasing-
upward parasequence (BDUP): a case study from
thè Lower Jurassic of thè Trento carbonate platform
(southern Alps), Italy. Rivista Italiana di Paleontolo¬
gia e Stratigrafia, Milano, 110 (1): 77-85.
Giannetti A., Monaco P., Caracuel J. E., Soria J. &
Yébenes A., 2007 - Functional morphology and
ethology of decapod crustaceans gathered by Tha¬
lassinoides branched burrows in Mesozoic shallow
water environments. In: 3rd Symposium on Mesozoic
and Cenozoic Decapod Crustaceans. Museo di Storia
Naturale di Milano, May 23-25, 2007. A. Garassino,
R. M. Feldmann & G. Teruzzi (eds.). Memorie della
Società italiana di Scienze naturali e del Museo civico
di Storia naturale di Milano, Milano, XXXV (II): 48-
52.
Hasiotis S. T. & Mitchell C. E., 1993 - A comparison of
crayfìsh burrow morphologies: Triassic and Holocene
fossil, paleo- and neo-ichnological evidence, and thè
identification of their burrowing signatures. Ichnos, 2:
291-314.
Howard J. D. & Frey R. W., 1984 - Characteristic trace
fossils in nearshore to offshore sequences, Upper Cre¬
taceous of east-central Utah. Canadian Journal Earth
Science, 21: 200-219.
Ksiqzkiewicz M., 1970 - Observations on thè ichno-
fauna of thè Polish Carpathians. In: Trace Fossils. T.
P. Crimes & J. C. Harper (eds.). Geological Journal
Special lssue, Liverpool.
Ksiqzkiewicz M., 1977 - Trace fossils in thè flysch of thè
Polish Carpathians. Paleontologica Polonica, War-
zawa, 36: 1-208.
.
82
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Lundgren B., 1891 - Studier òfver fossilfòrande Iòsa
block. Geoliska Fòreningens Stockholm Fòrhandlin-
gar, Stockholm, 13: 111-121.
Martinsson A., 1970 - Toponomy of trace fossils. In:
Trace Fossils. T. P. Crimes & J. C. Harper (eds.). Geo¬
logico il Journal, Special Issue.
Monaco P., 2000 - Decapod burrows ( Thalassinoides ,
Ophiomorpha ) and crustacean remains in thè Calcari
Grigi, lower Jurassic, Trento platform (Italy). 1° work¬
shop on Mesozoic and Tertiary decapod crustaceans.
Studi e Ricerche - Associazione Amici Museo - Museo
Civico “G. Zannato ”, Montecchio Maggiore (VI), oct.
4-6,2000:55-57.
Monaco P., in press - Taphonomic features of Paleodic-
tyon and other graphoglyptid trace fossils in Oligo-
Miocene thin-bedded turbidites of Northern Apen-
nines flysch deposits (Italy). Palaios.
Monaco P. & Garassino A., 2001 - Burrows and body
fossil of decapod crustaceans in thè Calcari Grigi,
Lower Jurassic, Trento platform (Italy). Geobios,
Lyon, 34 (3): 291-301.
Monaco P. & Giannetti A., 2002 - Three-dimensional
burrow systems and taphofacies in shallowing-upward
parasequences, lower Jurassic carbonate platform (Cal¬
cari Grigi, Southern Alps, Italy). Facies, 47: 57-82.
Monaco P., Giannetti A., Caracuel J. E. & Yébenes A.,
2005 - Lower Cretaceous (Albian) shell-armoured
and associated echinoid trace fossils from thè Sàcaras
Formation, Serra Gelada area, southeast Spain.
Lethaia, Oslo, 38: 1-13.
Monaco P., Nocchi M., Ortega-Huertas M., Palomo
I., Martinez F. & Chiavini G., 1994 - Depositional
trends in thè Valdorbia section (Central Italy) during
thè Early Jurassic, as revealed by micropaleontology,
sedimentology and geochemistry. Eclogae geologicae
Helvetiae, 87 (1): 157-223.
Palmer T. J., 1978 - Burrows at certain omission surfaces
on thè Middle Ordovician of thè Upper Mississippi
Valley. Journal of Paleontology, Lawrence, 52: 109-
117.
Rieth A., 1 932 - Neue Funde spongeliomorpher Fucoiden
aus dem Jura Schwabens. Geologische und Palàon-
tologische Abhandlungen,ÌA.¥. 19: 257-294.
Seilacher A., 1964- Biogenic sedimentary structures. In:
Approaches to Paleoecology. J. Imbrie & N. Newell
(eds.). Wiley, New York.
Seilacher A., 1977 - Pattern analysis of Paleodictyon and
related trace fossils. In: Trace Fossils 2. T. P. Crimes &
J. C. Harper (eds.). Geologica! Journal, Special Issue
9, London.
Sheehan P. M. & Schiefelbein J. D. R., 1984 - The trace
fossil Thalassinoides from thè Upper Ordovician of
thè eastem Great Basin, deep burrowing in thè Early
Paleozoic. Journal of Paleontology, Lawrence, 58:
440-447.
Tchoumatchenco P. & Uchman A., 2001 - The oldest
deep-sea Ophiomorpha and Scolicia and associated
trace fossils from thè Upper Jurassic-Lower Creta¬
ceous deep-water turbidite deposits of SW Bulgaria.
Palaeogeography, Palaeoclimatology, Palaeoecolo-
gy, Amsterdam, 169: 85-99.
Tunis G. & Uchman A., 2003 - Trace fossils from thè
Brkini flysch (Eocene), south-westem Slovenia. Gor-
tania - Atti del Museo Friulano di Storia Naturale,
Udine, 25: 31-45.
Uchman A., 1995 - Taxonomy and paleoecology of
flysch trace fossils: thè Mamoso-arenacea Formation 5
and associated facies (Miocene, Northern Apennines,
Italy). Beringeria, Heft, 15: 1- 116. c
Uchman A., 1998 - Taxonomy and ethology of flysch li
trace fossils: revision of thè Marian Ksiqzkiewicz
collection and studies of complementary material.
Annales So-cietatis Geologorum Poloniae, Kraków,
68: 105-218.
Uchman A. & Demircan H., 1999 - Trace fossils of
Miocene deep-sea fan fringe deposits from thè Cingoz
Formation, southern Turkey. Annales Societatis Geo¬
logorum Poloniae, Warzawa, 69: 125-135.
1 11
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Paolo Monaco - Dipartimento di Scienze della Terra, Università di Perugia, piazza dell’Università, 06100 Perugia, Italy.
e-mail: pmonaco@unipg.it
Jesus E. Caracuel - Dpto. Ciencias de la Tierra y del Medio Ambiente, Universidad de Alicante, Apdo. 99, 03080 Alicante, Spain.
e-mail: jesus.caracuel@ua.es
Jesus M. Soria - Dpto. Ciencias de la Tierra y del Medio Ambiente, Universidad de Alicante, Apdo. 99, 03080 Alicante, Spain.
e-mail: jesus.soria@ua.es
Alfonso Yébenes - Dpto. Ciencias de la Tierra y del Medio Ambiente, Universidad de Alicante, Apdo. 99, 03080 Alicante, Spain.
e-mail: ayeb@telefonica.net
Alice Giannetti - Geologisches Institut, Bonn Universitat, NuBallee 8, 531 15 Bonn, Germany.
e-mail: giannetti@geo.uni-bonn.de
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Carlos Neto de Carvalho & Pedro Andrade E. J. Viegas
Microfacies analysis of Thalassinoides infìlling:
causes of collective burial among Mecochirus populations
Decapod ichnofossils are thè commonest ones in thè
stratigraphic record of Portugal. Among them, Thalassi¬
noides dominates thè lagoonal and inner shelf facies
from thè late Sinemurian (Duarte et al. 2006) to Messin-
ian (Cachào et al. 2000) of west Iberia defining, many
times, thè entire carbonate sequence with its “nodular”
fabric. However, as usuai all over thè world, thè deca¬
pod producer and their burrows almost never are found
preserved together, allowing identification of thè decapod
modus vivendi. Recently, Neto de Carvalho et al. (2007)
described an unit with burrow mazes of Thalassinoides
suevicus (Rieth, 1932) with hundreds of Mecochirus
rapax (Harbort, 1905) as an ohrution lagerstàtte, in thè
lower Barremian from thè Lusitanian Basin. Those lob-
sters show an exceptionally complete preservation of thè
thin carapace including thè first pereiopods reaching 12
cm, as well as all thè omamentation details of thè cepha-
lotorax and pereiopods. Mecochirus are usually enrolled
in thè rigor mortis position (Fig. la) and are laterally
deposited dose to thè floor of thè burrows. The environ-
mental causes that lead to thè collective and instantane-
ous entombment of Mecochirus populations within their
burrow systems, repeated several times during as much as
1 Ma (Rey, 1993), apparently never to happen again in all
thè Lusitanian basin is unknown.
The Mecochirus-Thalassinoides association occurs
in a Lower Cretaceous sequence of mixed sedimentary
shelf, carbonate and siliciclastic sediments (Rey, 1993).
Fossils crop out, along more than 700 m on a monoclinal
1 5° north-dipping sequence, without evidence of impor¬
tai tectonic or strong diagenetic imprint. Assigned to
thè top of thè lower Barremian (Rey, 1972), thè Boca do
Chapim Formation is composed of marly limestones that
are dominated in thè first meters by poorly sorted, suban-
gular quartz grains, less than 4 cm, and bioclastic at thè
top, with subordinate intercalations of argillaceous marls.
The Thalassinoides ichnoguild and benthic fauna found
in thè Boca do Chapim Formation indicate a back-reef
lagoon in a shallow subtidal setting (cf. Rey, 1972; Neto
de Carvalho et al. 2007), with episodio coarse influxes
from a proximal fluvial braided System (Fig. lb). In fact,
to thè east, thè Boca do Chapim Formation interfingers
with a siliciclastic complex of fluvial origin. Microfa¬
cies analysis of Thalassinoides fili permits description of
thè paleoenvironmental events responsible for death and
complete preservation of Mecochirus without evidences
of scavenging or decay. Thin section analysis revealed
4 microfacies, all defined by thè absence of abundant
organic remains (wood fragments) or pyrite, indicators of
anoxia. Curiously, Favreina was not reported which could
be thè result of a predatory feeding habit for thè producer
and a frequent interaction with thè marine bottoni. Micro¬
facies #1 is composed of biomicrite with abundant shell
fragments and dasycladaceans. On thè other hand, micro¬
facies #2 is formed by intramicrite (Fig. le). Both micro¬
facies have little fine grained siliciclastics and almost no
Choffatella. These are evidences for a rapid infìlling of
Thalassinoides with dense mud and pelleted mud flows
from thè surroundings during storm events or strong tidal
currents. Biomicrites with Choffatella from thè intermedi-
ary microfacies #3 (Fig. ld), show more influence from
Coastal sediments. On thè other hand, microfacies #4,
composed of biomicrites, wackestones, and intrabiomi-
crosparites packstones (Fig. le), reveals abundant het-
erogranular siliciclastics (abundant quartz and quartzite,
angular to sub-rounded grains not more than 5 mm long).
This microfacies indicates episodic flows of coarse sedi-
ment from thè coast and thè fluvial braided System (with
contributions from alluvial fans) nearby thè lagoon sector
(Fig. 1 f), leading to successive smothering of Mecochi¬
rus by thè flood runoff with sudden changes in thè water
chemistry (<salinity, <pH) within Thalassinoides.
References
Cachào M., de Gibert J. M., Mayoral E., Muniz F. & da Silva
C. M., 2000 - Paleoicnologia da Forma9ào de Cacela
(Miocénio Superior), Algarve, Portugal: dados prelimin-
ares. I Congresso Ibèrico de Paleontologia/XVI Jomadas
de la Sociedad Espahola de Paleontologia, Évora: 5-7.
Duarte L. V., Perilli N., Antonioli L., Rodrigues R., Cabrai
M. C., Dino R. & Azerédo A. C., 2006 - Evidèncias
sedimentológicas, geoquimicas (COT) e micropaleon-
tológicas nas fàcies betuminosas do Sinemuriano ter¬
minal de Agua de Madeiros (Portugal). J. Mirào & A.
Balbino (eds.). VII Congresso Nacional de Geologia,
Livro de Resumos, Évora, 2: 633-636.
Neto de Carvalho C., Viegas, P. A. & Cachào, M., 2007 -
Thalassinoides and its Producer: Populations of Meco¬
chirus Buried within their Burrow Systems, Boca do
Chapim Formation (Lower Cretaceous), Portugal.
Palaios, 22: 107-112.
Rey J., 1972 - Recherches géologiques sur le Crétacé
Inférieur de L’ Estremadura (Portugal). Memórias dos
Servigos Geológicos de Portugal, 21 : 1-477.
Rey J., 1993 - Stratigraphie séquentielle sur une piate-
forme à sédimentation mixte: exemple du Crétacé
inférieur du Bassin Lusitanien. Comunicagòes dos
Servigos Geológicos de Portugal, 79: 87-97.
84
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Fig. 1 - Thin sections analysis of thè Thalassinoides suevicus fili that shrouded Mecochirus rapax in thè obrutionary deposits from
Boca do Chapim Formation. a) Mecochirus in rigore mortis showing details of omamentation; coin = 24 mm. b) Coarse quartzarenite
fili of Thalassinoides suevicus in brackish-water intertidal sandy bars; coin = 22 mm. c) Thalassinoides fili of microfacies #2 type
(intramicrosparite), with large muddy clasts and Miliolidae. Compare with thè micritic matrix (top right). d) Biomicrite with Chof-
fatella (microfacies #3) and thè surrounding matrix (mudstone). Note thè lack of burrow wall. e) Microfacies #4: Intrabiomicrosparite
packstone with sub-rounded quartzite grains showing a metamorphic (Variscan) source for thè siliciclastics. Bar scale = 0,5 mm.
f) Paleogeographic reconstitution of thè studied area during thè Lower Cretaceous. Artwork of Nuno Dias and Carlos Farinha.
Carlos Neto De Carvalho - Geology and Paleontology Office, Geopark Naturtejo Meseta Meridional - UNESCO European and Global
Geopark. Avenida Zona Nova de Expansào, 6060-101 Idanha-a-Nova, Portugal.
e-mail: paleo@walla.com
Pedro Andrade E. J. Viegas - Rua Olivai Santo, Lote 1, 3° Esquerdo, 2625-585 Vialonga, Portugal.
e-mail: paleomail@gmail.com
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Cristina M. Robins, Rodney M. Feldmann & Carrie E. Schweitzer
Primitive brachyurans and galatheids from Ernstbrunn, Austria:
an evaluation of thè Friedrich Bachmayer Collection
The Friedrich Bachmayer Collection, housed in thè
Naturai History Museum of Vienna, Austria, contains
many specimens of primitive brachyurans and galatheids
ffom thè Upper Jurassic (Tithonian) of Ernstbrunn, Austria.
Over 130 specimens, a small fraction of his extensive col¬
lection, were selected and studied based on their excellent
preservation and unique characteristics. Special effort was
made to choose specimens that contained appendages, par-
ticularly chelae, or possible ventral carapace surfaces.
The geology of thè Ernstbrunn area has been extensively
studied for both paleontological and tectonic reasons (Zeiss,
2001; Adàmek, 2005). The Ernstbrunn Limestone reaches
120 m in thickness (Adàmek, 2005) and is sandwiched
between thè Waschberg Zone and thè Molasse Basin
(Zeiss, 2001). Stratigraphically, thè Ernstbrunn Limestone
is unconformably below thè Klement Formation and con-
formably overlies thè Klentnice Formation (Zeiss, 2001).
i It crops out in several locations in northem Austria and thè
Czech Republic (Zeiss, 2001; Adàmek, 2005). It caps thè
Malmian sedimentation sequence, a basinal pelagic-carbon-
ate to marginai carbonate sequence that was most likely
deposited during thè Tithonian (Adàmek, 2005). In addition
to crustaceans, fossils found in thè Ernstbrunn Limestone
include foraminifera, green algae, sponges, hydrozoans,
corals, cephalopods, bryozoans, brachiopods, mollusks, and
echinoderms, isopods as well as fish (Zeiss, 2001; Adàmek,
2005). Given thè presence of both coral and algal fossils,
thè Ernstbrunn Limestone was deposited within thè photic
zone. Bachmayer (1947) hypothesized that thè crustaceans
he studied lived on a reef talus slope. Zeiss (2001) agreed
in pari, but offered a hypothesis of a lagoonal setting that
opened up into a marine reef environment.
During thè lower Miocene thè Carpathian orogeny
caused tilting and uplift within this area (Zeiss, 2001;
Adàmek, 2005). Subsequent tectonic events have buried
much of thè Jurassic units in this area undemeath a thick
series of flysch deposits. Thus, outcrops are limited and
much of thè information about thè Ernstbrunn Limestone
was taken from drill-core samples (Adàmek, 2005).
The Ernstbrunn Limestone has been quarried for use
as a building rock; also, since it is so pure, it is used for
making plaster. Castle Staatz and Castle Falkenstein (now
ruins) were constructed from thè Ernstbrunn Limestone
(Zeiss, 2001).
All but two of thè selected decapod specimens are pre-
served within white, fine-grained crystalline limestone.
Bachmayer (1947) described several localities where he
found crustacean specimens, including an abandoned
quarry near Ernstbrunn, Austria, which contained white,
fine-grained limestone that yielded many crustacean
fossils. It appears that this quarry is stili accessible. It is
possible thè fossils are from this locality, as it is thè only
one he described that is both fossil-rich and of white lime¬
stone. However, Bachmayer also noted that whereas most
of thè specimens found at Ernstbrunn were very similar or
identical to those found in Stramberg, now in thè Czech
Republic, thè ones from Ernstbrunn were barely half thè
size of thè ones from Stramberg. Given thè relatively large
size of thè individuate within this collection, it is possible
that some of thè fossils originated from Stramberg, Czech
Republic, and are mixed in with thè Ernstbrunn fossils.
The majority of thè studied specimens are from thè
superfamily Galatheidea Samouelle, 1819, and primitive
brachyurans that are members of thè families Prosopidae
von Meyer, 1860, and Goniodromitidae Beurlen, 1932
(synonym Pithonotidae Glaessner, 1933), both of which
are considered key to understanding thè evolution of
brachyurans (Muller et al., 2000; Feldmann et al., 2006).
In many specimens, all that remains of thè dorsal carapace
is a thin chalky layer; however, fine details of omamenta-
tion and other morphological features are visible. Unfor-
tunately, similar to other known Jurassic brachyurans,
none of thè specimens is articulated. While there are
numerous disarticulated appendages and chelae among
thè specimens, no definitive associations between chelae
and individuai species can be made.
The comparatively excellent preservation and variety of
these specimens allows detailed study of decapods that are not
widely documented within thè fossil record. Based on thè high
quality of preservation and conclusions from previous work
on thè paleogeography and paleoenvironment of thè Late
Jurassic (Bachmayer, 1947; Muller et al, 2000; Feldmann et
al., 2006), it is concluded that thè specimens were preserved in
a coral-dominant reef setting typical of thè Tithonian.
References
Adàmek J., 2005 - The Jurassic floor of thè Bohemian
Massif in Moravia-geology and paleogeography. Bul-
letin of Geosciences, Czech Geological Survey, Bmo,
80: 291-305.
Bachmayer F., 1947 - Die Crustaceen aus dem Ernstbrun-
ner Kalk der Jura-Klippenzone zwischen Donau und
Thaya. Jahrbuch der Geologischen Bundesanstalt,
Vienna, 8: 35-47.
Beurlen K., 1932 - Brachyurenreste aus dem Lias von
Bomholm mit Beitràgen zur Phylogenie und Syste-
matik der Brachyuren Dekapoden. Palàontologische
Zeitschrift, Stuttgart. 14: 52-66.
86
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Feldmann R. M., Lazàr I. & Schweitzer C. E., 2006 -
New crabs (Decapoda: Brachyura: Prosopidae) from
Jurassic (Oxfordian) sponge bioherms of Dobrogea,
Romania. Bulletin of thè Mizunami Fossil Museum,
Mizunami, Japan, 33: 1-20.
Glaessner M. F., 1933 - Die Krabben der Jura-Formation.
Zentralblatt fur Mineralogie, Geologie und Palaonto-
logie, Stuttgart. Abt. B: 178-191.
Miiller R, Krobicki M. & Wehner G., 2000 - Jurassic and
Cretaceous primitive crabs of thè Family Prosopidae
(Decapoda: Brachyura). Their taxonomy, ecology and
biogeography. Annales Societatis Geologorum Polo-
niae, Kraków, 70: 49-79.
Meyer H. von, 1860 - Die Prosoponiden oder die Familie
der Maskenkrebse. Palaeontographica, Stuttgart, 7,
183-222.
Samouelle G., 1819 - The Entomologist’s Useful Com-
pendium, or An Introduction to thè Knowledge of
British Insects. Thomas Boys, London.
Zeiss A., 2001 - Die Ammonitenfauna der Tithonklippen
von Emstbrunn, Niederòsterreich. Neue Denkschrif-
ten des Naturhistorischen Museums in Wien , Vienna:
14-26.
Cristina M. Robins - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: crobins@kent.edu
Rodney M. Feldmann - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rfeldman@kent.edu
Carrie E. Schweitzer - Dept. of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, U.S.A.
e-mail: cschweit@kent.edu
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Giinter Schweigert
Preservation of decapod crustaceans in thè Upper Jurassic
lithographic limestones of southern Germany
The lithographic limestones in thè Upper Jurassic of
southern Germany belong to thè most important Fossil-
lagerstàtten conceming not only vertebrates but also thè
fossil record of decapod crustaceans in thè Jurassic and
even in thè whole Mesozoic. There are numerous sites in
this region, often generalized and labelled as “Solnhofen”,
but in fact coming from various localities (e.g. Brunn,
Daiting, Eichstàtt, Hienheim, Langenaltheim, Mòm-
sheim, Nusplingen, Painten, Pfalzpaint, Schamhaupten,
Solnhofen, Wattendorf, Zandt, Fig. 1 A) in a large area and
spanning thè time interval from thè Late Kimmeridgian to
thè Early Tithonian (Schweigert & Garassino, 2003; Ga-
rassino & Schweigert, 2006; Schweigert, in press). Only
very few of these localities did not yield any crustaceans,
possibly due to unfavourable preservation conditions
during earliest diagenesis.
Formerly thè focus of thè scientific interest lay almost
exclusively in thè description of taxa and was stili very
successful in thè last decade because of several scien¬
tific excavations (Brunn, Nusplingen, Schamhaupten,
Wattendorf), intensive evaluation of older collections,
enhanced quality of preparation and new preparation
techniques, and thè use of ultraviolet illumination for
detailed anatomical studies (Polz, 1999, 2000; Garassino
& Schweigert, 2006; Schweigert 200 la, b, c, 2002, 2003;
Schweigert et al., 2000; Schweigert & Garassino, 2004,
2005a, b, 2006). Palaeoecological or functional analyses
of decapod crustaceans from thè lithographic limestones
have been mostly neglected in thè past. To date c. 85 valid
taxa have been reported - younger synonyms, juveniles
or sexual dimorphs not included. Stili three to four com-
pletely new taxa appear from thè quarries every year.
However, this high number is summarized from thè vari¬
ous localities which have different lithologies and ages. In
thè distribution of taxa significant locai differences exist,
probably due to different ecologies in thè surroundings.
At present we try to reconstruct thè former habitats
in which decapods lived and thè circumstances of their
burial. The bulk of crustacean fossils, lobsters as well as
shrimps and prawns, are not true “body fossils” but exu-
viae. In most cases thè exuviae can be easily recognized
by their strong compaction, whereas in body fossils there
is stili a remarkable relief and often a phosphatic pres¬
ervation of thè soft tissue present (Briggs et al., 2005).
Sometimes exuviae also differ significantly from “body
fossils” in their orientation. Moreover, exuviae often show
typical disaggregating, like incomplete articles, a dorsally
opened carapace and, after longer time of drifting, numer¬
ous nematode borings. A common preservational state of
many eryonids shows their pereiopods I pointing back-
ward. This was caused by anchoring of thè heavy chelae
hanging down from thè drifting exuvia (Fig. 2A). During
decay thè carapace of eryonids splits off in thè dorso-
median line suggesting a much broader carapace than in
reality. In thè past this phenomenon has led to erroneous
reconstructions and/or thè erection of synonymous taxa.
The co-occurrence of different ontogenetic stages of a
single palaeobiospecies suggests an identity of thè habitat
and thè place of burial, or at least a closest neighbourhood.
In thè lithographic limestones in thè vicinity of Eichstàtt
this is true for several shrimps and prawns ( Aeger Miin-
ster, 1839, Antrimpos Miinster, 1839, Hefriga Miinster,
1839). Most other taxa, however, are very rare or even
only known by unique specimens (Schweigert, 2002;
Schweigert & Garassino, 2004, 2005a, b). They also
mostly represent exuviae. For this reason it is assumed
that they must have lived in more distant areas, and
their record often depended on random circumstances.
More data about their former habitats and life styles are
available from a functional analysis, e.g. of thè chelae.
Among shrimps there are typical detritus feeders with
pincer-like chelae (e.g. Blaculla Miinster, 1839, Dusa
Miinster, 1839) which must have lived in reefal habitats.
Brachyurans (prosopids, galatheids) often occur in Upper
Jurassic sponge-microbial reefs or adjacent bedded lime¬
stones (Miiller et al., 2000), whereas they are extremely
rare within thè lithographic limestones (e.g. Òchselberg
near Breitenhill, Garassino et al., 2005). Anomurans
(e.g. Maglia Miinster, 1839) are common in calcare-
ous, bioturbated deposits, but also very rarely recorded
from lithographic limestones. Interestingly, some of thè
Late Jurassic anomuran taxa are only recorded fforn thè
lithographic limestones, and not from coeval bioturbated
basin deposits (e.g. Etallonia Oppel, 1861, Megachela
Schweigert, 2003). Possibly anomurans are often over-
looked due to their poor sclerotization and preservation
and small size (Polz, 1999; Schweigert, 2003).
Endobenthic decapods like glypheids occur frequently
(Polz, 2000; Schweigert & Garassino, 2005a, 2006), but
almost exclusively as exuviae. Notably there are some
locai occurrences, like that of Glyphea saemanni Oppel
within thè lithographic limestone of Brunn in eastem
Bavaria (Ròper et al., 1996). The latter was originally
described from thè famous locality of Cerin (Dpt. Ain,
France), thus pointing to a special ecological niche being
present in both localities. The thalassinoid burrows of
glypheids ( Spongeliomorpha ) are recorded from only a
few localities (Brunn, Nusplingen) but yet without in-situ
findings.
Epibenthic lobsters which exhibit strong chelae, like
erymids ( Eryma v. Meyer, 1840, Erymastacus Beurlen,
1928, Palaeastacus Bell, 1850, Pustulina Quenstdt,
1857), are significantly more common in localities in
which also other benthic organisms occur (e.g. Zandt).
88
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
This is especially thè case conceming adult specimens.
An autochthonous occurrence can be proved by thè
record of interactions between thè animals and thè sub-
strate. A good example is thè prawn Antrimpos unde-
narius Schweigert, 2001 , which is thè most frequent deca-
pod in thè small but rather deep lagoon of Nusplingen in
thè western part of thè Swabian Alb (Fig. 1 A; Schweigert,
200 lb). Besides exuviae there are plenty of specimens
deadly injured by predators (c. 50 % of all specimens),
and often they left their arrow-shaped last traces on thè
seafloor ( Telsonichnus Schweigert, 1998), produced by
thè tail fan (Schweigert, 1998, 200 lb; Schweigert & Dietl,
2005). Mass occurrences of small, comma-shaped copro-
lites with plant debris might also stem from this prawn
that obviously lived in great abundance in this lagoon.
Remains of Antrimpos undenarius Schweigert, 2001,
are recorded as thè prey of nautilids, fìshes, and teuthoid
squids. Interestingly, in thè Nusplingen Lithographic
Limestone thè juveniles or larva of Antrimpos are lacking.
Most likely they did not live in thè lagoon but somewhere
else. Such juvenile shrimps and prawns are quite fre¬
quent on some bedding planes in thè Eichstàtt area (e.g.
“ Bombur complicatus ” by Miinster, 1839). Resting traces
of palinurids ( Tripartichnus Vallon & Ròper, 2006) occur
in thè lithographic limestones of Walting near Pfalzpaint,
whereas thè animals themselves are lacking there (Vallon
& Ròper, 2006).
A passive input of crustacean remains (e.g. isolated
chelae) by predators is important in small lagoons like
that of Nusplingen (Schweigert et al., 2000). But there
are even some rare examples of benthic decapods prob-
ably sunken down from drifting wood to thè seafloor
(Fig. 1B). Several recent findings of thè common reptant
Mecochirus longimanatus (Schlotheim, 1820) from thè
Solnhofen Lithographic Limestone of Langenaltheim are
spectacular. They show thè act of moulting preserved on
thè bedding piane (Fig. 2B) and some examples will be
described in detail elsewhere. Because of thè generai of
lack of bioturbation, thè lithographic limestones are often
said to have been deposited in a restricted environment,
uncomfortable for higher organisms. For thè main rea-
sons, either oxygen depletion or hypersaline brines are
discussed. However, for several specialized taxa, like
Mecochirus Germar, 1927, life conditions on thè sea floor
were tolerable at least temporarily. Some of thè benthic
eryonids show adaptations to soft substrates, especially in
Eryon arctiformis (Schlotheim, 1820). Similar to Meco¬
chirus (cf. Oppel, 1862, PI. 23, figs. 1-2) its carapace and
parts of thè appendages exhibit a broad fringed margin
composed of fine setae. This could either be interpreted as
a “snowshoe effect” hampering thè sinking into thè soft
ground, or thè respiration was enhanced by thè enormous
enlargement of thè body surface thus enabling thè dwell-
ing in somewhat low-oxygen environments. The fringed
margin is only preserved in specimens which were imme-
diately buried during rapid sedimentation events, but
usually not in exuviae. Òther eryonids (e.g. Cycleryon
Glaessner, 1965) exhibit large stalked eyes pointing to
coarser substrates which allowed thè animals a shallow
trenching, waiting for prey.
Also in thè very rare caridean shrimp Pleopteryx
kuempeli Schweigert & Garassino, 2004, an adapta-
tion to low-oxygen conditions was recently suggested,
because of enigmatic, very large feather-like pleopodal
appendages developed, unknown from any Recent genus
(Schweigert & Garassino, 2004, 2006). The relative
abundance of this taxon in a small area in thè northern
vicinity of Eichstàtt (Wintershof) points to a dose dis-
tance from thè place of burial to thè originai habitat of
this shrimp.
Summarizing thè above observations, it can be stated
that there are only a few specialized crustaceans living
in thè same area in which thè lithographic limestones
were deposited. However, thè bulk of decapod crustacean
taxa recorded from thè lithographic limestones must be
regarded as allochthonous.
Fig. 1 - A) Sites of lithographic limestones in southern Germany which contain decapod crustaceans (from Fiirsich et al., in press,
slightly modified). B) Eryon arctiformis (Schlotheim, 1820). Specimen probably sunken from drifting wood to thè seafloor, with thè
ventral side upwards. Solnhofen Lithographic Limestone, Langenaltheim near Solnhofen, Hybonotum Zone; Jura-Museum Eichstàtt,
no. JME-SOS 6816a. Length of specimen c. 82 mrn.
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
89
Fig. 2 - A) Cycleryon propinquus (Schlotheim, 1822) (= type specimen of Eryon rehmanni v. Meyer, 1838). Solnhofen Lithographic
Limestone, vicinity of Solnhofen; Lower Tithonian, Hybonotum Zone; Furstlich-Furstenbergische Sammlung Donaueschingen, no.
Pz 2799.1. Length of specimen c. 170 mm. B) Mecochirus longimanatus (Schlotheim, 1820), exuvia lacking thè carapace, with
traces of thè moulting. Solnhofen Lithographic Limestone, Langenaltheim near Solnhofen, Hybonotum Zone; Staatliches Museum fìir
Naturkunde Stuttgart, no. 23262. Length of specimen c. 60 mm.
References
Briggs D. E. G., Moore R., Shultz J. W. & Schweigert G.,
2005 - Mineralization of soft-part anatomy and invad-
ing microbes in thè horseshoe crab Mesolimulus from
thè Upper Jurassic Lagerstàtte of Nusplingen, Ger-
many. Proceedings of thè Royal Society of London ,
series B, London, 272: 627-632.
Fiirsich F. T., Màuser M., Schneider S. & Werner W.,
in press - The Wattendorf Plattenkalk (Upper Kim-
meridgian) - a new conservation lagerstàtte from thè
northem Franconian Alb, southern Germany. Neues
Jahrbuch fur Geologie und Palàontologie Abhandlun-
gen, Stuttgart.
Garassino A., De Angeli A. & Schweigert G., 2005 -
Brachyurans from thè Upper Jurassic (Kim-
meridgian - Tithonian) lithographic limestones of
Pfalzpaint and Breitenhill (Bavaria, S Germany).
Atti della Società italiana di Scienze naturali e del
Museo civico di Storia naturale in Milano , Milano,
146 (1): 69-78.
Garassino A. & Schweigert G., 2006 - The Upper
Jurassic Solnhofen decapod crustacean fauna: review
of thè types from old descriptions. Part I. lnfraorders
Astacidea, Thalassinidea, and Palinura. Memorie della
Società italiana di Scienze naturali e del Museo civico
di Storia naturale di Milano, Milano, 34 (1): 1-64.
Muller P., Krobicki M. & Wehner, G., 2000 - Jurassic and
Cretaceous primitive crabs of thè family Prosopidae
(Decapoda: Brachyura) - their taxonomy, ecology and
biogeography. Annales Societatis Geologorum Polo-
niae, Kraków, 70: 49-79.
Miinster, G. Graf zu, 1839 - Decapoda Macroura. Abbil-
dung und Beschreibung der fossilen langschwànzigen
Krebse in den Kalkschiefem von Bayem. Beitràge zur
Petrefactenkunde, Bayreuth, 2: 1-88.
Oppel A., 1862 - Ùber jurassische Crustaceen. Palaeon-
tologìsche Mittheilungen aus dem Museum des kònig-
lich Bayerischen Staates, Stuttgart, 1: 1-120.
Polz H., 1999 - Etal Ionia hoellorum sp. nov. (Crustacea,
Decapoda, Axiidae) aus dem Oberkimmeridgium der
sudlichen Frankenalb. Archaeopteryx, Eichstàtt, 17:
33-39.
Polz H., 2000 - Glyphea viohli sp. nov. (Crustacea: Deca¬
poda: Glypheidae) aus den Solnhofener Plattenkalken.
Archaeopteryx, Eichstàtt, 18: 17-25.
Ròper M., Rothgaenger M. & Rothgaenger K., 1996
Die Plattenkalke von Brunn, Landkreis Regensburg.
Eichendorf (Eichendorf- Ver lag).
Schweigert G., 1998 - Die Spurenfauna des Nusplinger
Plattenkalks (Oberjura, Schwàbische Alb). Stuttgarter
Beitràge zur Naturkunde, Serie B, Stuttgart, 262: 1-47.
90
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Kimmeridgium, Schwabische Alb) im Vergleich mit
frànkischen Vorkommen. Stuttgarter Beitràge zur
Naturkunde, Serie B, Stuttgart, 285: 1-25.
Schweigert G. & Garassino A., 2003 - New studies of
decapod crustaceans from thè Upper Jurassic litho-
graphic limestones of southern Germany. Contributions
to Zoology, The Hague, 72: 173-179.
Schweigert G. & Garassino A., 2004 - New genera and
species of shrimps (Crustacea: Decapoda: Dendro-
branchiata, Caridea) from thè Upper Jurassic litho-
graphic limestones of S Germany. Stuttgarter Beitràge
zur Naturkunde, Serie B, Stuttgart, 350: 1-33.
Schweigert G. & Garassino A., 2005a - The lobster genus
Squamosoglyphea Beurlen, 1930 (Crustacea: Deca¬
poda: Glypheidae) in thè Upper Jurassic lithographic
limestones of southern Germany. Neues Jahrbuch fur
Geologie und Palàontologie Monatshefte, Stuttgart,
2005: 269-288.
Schweigert G. & Garassino A., 2005b - First record of thè
shrimp genus Carpopenaeus Glaessner, 1945 (Crus¬
tacea: Decapoda: Carpopenaeidae) from thè Jurassic.
Neues Jahrbuch fur Geologie und Palàontologie
Monatshefte, Stuttgart, 2005: 490-502.
Schweigert G. & Garassino A., 2006 - News on Pleop-
teryx kuempeli Schweigert & Garassino, an enigmatic
shrimp (Crustacea: Decapoda: Caridea: Pleopteryxoi-
dea superfam. nov.) from thè Upper Jurassic of S Ger¬
many. Neues Jahrbuch fur Geologie und Palàontolo¬
gie Monatshefte, Stuttgart, 2006: 449-461.
Vallon L. H. & Ròper M., 2006 - Tripartichnus (n. igen.) -
A new trace fossil from thè Buntsandstein (Lower
Triassic) and from thè Solnhofen Lithographic Lime-
stone (Upper Jurassic). Palàontologische Zeitschrift,
Stuttgart, 80: 156-166.
Giinter Schweigert - Staatliches Museum fur Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany.
e-mail: schweigert.smns@naturkundemuseum-bw.de
—
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Schweigert G., 200 la - Dimorphismus bei Krebsen der
Gattung Cycleryon (Decapoda, Eryonidae) aus dem
Oberjura Siiddeutschlands. Stuttgarter Beitràge zur
Naturkunde, Serie B, Stuttgart, 305: 1-21.
Schweigert G., 200 lb - Eine neue Art der Gattung
Antrimpos Munster (Crustacea, Decapoda, Penaeidae)
aus dem Oberjura Siiddeutschlands. Stuttgarter Bei¬
tràge zur Naturkunde, Serie B, Stuttgart, 307: 1-33.
Schweigert G., 200 le - The late Jurassic decapod spe¬
cies Aeger tipularius (Schlotheim, 1822) (Crustacea:
Decapoda: Aegeridae). Stuttgarter Beitràge zur
Naturkunde, Serie B, Stuttgart, 309: 1-10.
Schweigert G., 2002 - Zwei neue Gamelen (Decapoda:
Dendrobranchiata, Eukyphida) aus oberjurassischen
Plattenkalken Siiddeutschlands. Stuttgarter Beitràge
zur Naturkunde, Serie B, Stuttgart, 323: 1-11.
Schweigert G., 2003 - Megachela frickhingeri n. g. n. sp.
(Crustacea: Decapoda: Thalassinidea: Axiidae) aus
dem Solnhofener Plattenkalk (Ober-Jura, Bayem).
Stuttgarter Beitràge zur Naturkunde, Serie B, Stutt¬
gart, 333: 1-12.
Schweigert G., in press - Ammonite biostratigraphy as a
tool for dating Upper Jurassic lithographic limestones
from South Germany - fìrst results and open ques-
tions. Neues Jahrbuch fur Geologie und Palàontolo¬
gie Abhandlungen, Stuttgart.
Schweigert G. & Dietl G., 2005 - Miscellanea aus dem
Nusplinger Plattenkalk (Ober-Kimmeridgium, Schwa¬
bische Alb) 6. Die Spurengattung Telsonichnus . Jahres-
berichte und Mitteilungen des Oberrheinischen geolo-
gischen Vereins, neue Folge, Stuttgart, 87: 431-438.
Schweigert G., Dietl G. & Ròper M., 2000 - Die Panzer-
krebse der Familie Erymidae van Straelen (Crustacea,
Decapoda) aus dem Nusplinger Plattenkalk (Ober-
Carrie E. Schweitzer, Rodney M. Feldmann, Aubrey M. Shirk &
Iuliana Lazàr
Differential diversity in Jurassic sponge-algal and
coralline communities, Romania
Detailed collection and evaluation of decapod faunas
from various reef settings in eastem and centrai Romania
demonstrate diversity differences and differential habi¬
tat preferences early in thè history of thè Anomura and
Brachyura.
Abundant and diverse decapod faunas have been
reported from three sets of coral-dominated reef localities
in thè vicinity of thè southeastem Carpathian Mountains
near thè towns of Sinaia and Bra$ov, Romania. The reef
localities near Sinaia are generally grouped together with
that near Moroeni in faunal counts (Patrulius, 1959, 1966).
The Sinaia localities consist of olistolith blocks situated
in thè mountains above thè town of Sinaia; many are no
longer accessible due to recent housing and infrastructure
development in thè area. The Moroeni locality was a large
olistolith block near a tuberculosis sanitorium in thè village
of Moroeni, south of Sinaia; that block is all but grown
over with vegetation. Patrulius collected these areas exten-
sively and reported as many as 38 decapod species from
20 genera from this set of localities (Patrulius, 1966). Our
collecting in these localities yielded signifìcantly fewer
taxa. Unfortunately, Patrulius ’s systematic work on these
decapods was never published, and it is diffìcult to com¬
pare thè number of species from our work with Patrulius’s
list without an evaluation of his material. In addition to thè
decapods, our collecting also yielded corals, brachiopods,
pelecypods, and echinoderms from these localities.
The decapods from thè locality at Purcàreni were also
preliminarily reported by Patrulius (1966) but were never
formally published. He reported as many as 18 species in
1 2 genera from thè Purcàreni olistolith, known locally as thè
“coral rock” and located east of Bra§ov. Collecting of that
olistolith during thè field seasons of 2004 and 2005 yielded 6
additional species in 5 additional genera (Shirk, 2006). Shirk
(2006) also reported several species each of brachiopods,
corals, pelecypods, and gastropods as well as ammonoids,
crinoids, sponges, and trace fossils from this locality.
Mufiu & Bàdàlutà (1971) reported a small decapod
fauna from a reef locality southeast of thè Carpathians in
eastem Romania that included 7 species in 6 genera. They
also reported corals, sponges, brachiopods, and pelecy¬
pods associated with thè decapods. Thus, it is clear that
thè coral reef faunas, both in terms of decapods and other
invertebrates, were generally diverse and robust.
Analysis of thè decapods collected from sponge-algal
megafacies in southeastem Romania (Central Dobrogea)
demonstrates that such of differing geometries appear
to have hosted slightly different, low-diversity decapod
faunas that lived within thè magafacies structures. Lay-
ered sponge biostromes at thè Gura Dobrogei, Romania,
locality include four species of brachyurans and several
indeterminate claw fragments. The identifiable genera,
representing thè Goniodromitinae Beurlen, 1932, and Pro-
sopinae von Meyer, 1 860, of thè Prosopidae von Meyer,
1 860, include Goniodromites Reuss, 1 859, and Cyclopro-
sopon Lòrenthey in Lòrenthey & Beurlen, 1929, of thè
Goniodromitinae, and a new taxon within thè Prosopinae.
Giant, cylindrical, sponge bioherms in thè Cheia Valley,
not far from thè Gura Dobrogei locality, yielded not only
species of Goniodromites and Cycloprosopon as well as
claw fragments, similar to thè Gura Dobrogei locality, but
also a species of Pithonoton von Meyer, 1842 (Feldmann
et al., 2006). In addition, one of thè fragmental specimens
appears to have strong affinities with Pithonoton insigne
von Meyer, 1857, a species known only from sponge
megafacies or laminated limestones (G. Schweigert,
personal commun.; Collins & Wierzbowski, 1985). The
cylindrical bioherm structures also yielded sponges, anne-
lid worms, brachiopods, and belemnites (Feldmann et al.,
2006). The prosopid genus collected from thè Gura Dobro¬
gei locality was not present in thè cylindrical reefs in thè
Cheia Valley. Both sets of localities received about equal
collecting effort, based upon thè size of outcrop, and thè
same collecting team worked at both sets of localities, so
it seems unlikely that there was a collecting bias between
thè sites. The presence of claw and appendage fragments
at both localities suggests that thè animals were living
within thè sponge reefs. Had thè decapod fragments been
carried into thè area by currents, thè tiny appendage and
claw fragments, and even thè tiny carapaces, would prob-
ably have been destroyed. In addition, thè decapods col¬
lected in association with thè cylindrical sponge bioherms
showed some level of habit partitioning, with some show-
ing preferences for thè bioherm itself, thè inter-bioherm
talus, and thè centrai core area of thè bioherm (Feldmann
et al., 2006). This specialization for different parts of thè
bioherm as well as slightly different faunas in bioherms of
differing geometries suggests that habitat partitioning was
off to an early and eamest start within thè Brachyura.
Of thè decapods collected from thè coral reef locali¬
ties, most were not collected from thè sponge localities.
Only three genera, Cycloprosopon, Goniodromites, and
Pithonoton, were collected from both sponge and coral
reef localities. The new prosopid genus and thè possible
specimen of Pithonoton insigne are thè only decapod taxa
collected only from sponge-algal habitats. Thus, it appears
that many more genera were specialized for coral reef habi¬
tats than for sponge habitats, perhaps due to thè shallower,
better oxygenated, better illuminated environment there. It
is also probable that, due to thè nature of thè wide variety
92
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
*
of morphology of corals, there are more niches and physi-
cal spaces for tiny crabs to hide, reproduce, and molt than
would be available in sponge-algal habitats, composed of
cylindrical, conical, or layered sponges.
The three genera Cycloprosopon, Pithonoton, and
Goniodromites, appear to have been generalists, able to
survive in coral and sponge-algal habitats. The eurytopic
adaptations of these three genera may account for their
repeated presence in species and generic lists of Jurassic
crabs; certainly, various species of Goniodromites as well as
P. marginatum von Meyer, 1 842, are some of thè most com-
monly encountered taxa in Jurassic decapod studies. More
often than not, these identifications seem to be consistent
with thè historic conception of these taxa, based upon our
survey of material in several museums over thè past two
summers. If these taxa were indeed generalists, they would
have been able to survive in many environments which may
in tum have allowed their survival for a long duration of
geologie time, which may also contribute to their common-
ality in Jurassic deposits across Europe. Detailed examina-
tion of specimens of these taxa is ongoing in an attempt to
determine which morphological features specifically may
have contributed to their broad adaptability.
When evaluating thè Romanian reef decapods at thè
family and subfamily level, it is notable that several
groups are absent from thè sponge-algal habitats. Family
level classification for thè purposes of this work largely
follows Patrulius (1966), whereas subfamily level clas¬
sification follows Glaessner (1969). We do not consider
Laeviprosopon Glaessner, 1933, to be a homolid as did
Patrulius (1966), because it clearly lacks lineae homoli-
cae. Notably absent from thè sponge-algal habitats is thè
Galatheidae Samouelle, 1819, which is well-represented
in all three of thè coral reef localities surveyed here. Also
almost completely lacking from thè sponge-algal habitats,
with thè exception of one genus and species, is thè Pro-
sopinae. The Goniodromitinae is well-represented in both
coral reef and sponge-algal habitats, suggesting that as for
thè two included genera, Goniodromites and Pithonoton ,
thè subfamily itself may have been broadly adapted and
eurytopic. The same may also have been true for thè early
Dynomenidae Ortmann, 1892, well-represented in coral
reef habitats and represented by one genus and species
within sponge-algal habitats. Investigation of thè evolu-
tionary pattems within these families and their relation-
ships with reefs of varying types is ongoing.
Table 1 - Decapoda reported from thè Sinaia and Moroeni localities. Data from Patrulius (1959, 1966).
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
93
Table 2 - Decapoda reported from thè Ghergheasa locality. Data from Mutiu & Bàdàlutà (1971).
Table 3 - Decapoda reported from thè Purcàreni locality. Data from Patrulius (1966) and Shirk (2006).
94
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
References
Bell T., 1 863 - A monograph of thè fossil malacostracous
Crustacea of Great Britain, Pt. II, Crustacea of thè Gault
and Greensand. Palaeontographical Society Monograph,
London.
Beurlen K., 1928 - Die fossilen Dromiaceen und ihre Stam-
mesgeschichte. Palàontologische Zeitschrift, Berlin, 10:
144-183.
Beurlen K., 1932 - Brachyurenreste aus dem Lias von Bom-
holm mit Beitràgen zur Phylogenie und Systematik der
Brachyuren Dekapoden. Palàontologische Zeitschrift,
Berlin, 14: 52-66.
Collins J. S. H. & Wierzbowski A., 1985 - Crabs ffom thè
Oxfordian sponge megafacies of Poland. Acta Geologica
Polonica, Warzawa, 35 (1-2): 73-88.
Étallon A., 1861 - Notes sur les Crustacés Jurassiques du
bassin du Jura. Mémoires de la Società de l ’Agriculture,
des Sciences et Lettres de la Haute Saóne, Vesoul, 9: 129-
171.
Feldmann R. M., Lazàr I. & Schweitzer C. E., 2006 - New
crabs (Decapoda: Brachyura: Prosopidae) ffom Jurassic
(Oxfordian) sponge bioherms of Dobrogea, Romania.
Bulletin of thè Mizunami Fossil Museum, Mizunami, 33:
1-20.
Gemmellaro G. G., 1870 - Studi paleontologici sulla fauna
del calzare a Terebratula janitor del Nord di Sicilia. Parte
I, Stabilimento Tipografico Lao, Palermo.
Glaessner M. F., 1933 - Die Krabben der Juraformation. Zen-
tralblatt fiir Mineralogie, Geologie und Palaontologie,
Stuttgart, Abtheilung B: 178-191.
Glaessner M. F., 1969 - Decapoda, p. R400-R533, R626-628.
In: Treatise on Invertebrate Paleontology. R. C. Moore
(ed.). Pt. R4 (2). Geological Society of America and Uni¬
versity of Kansas Press, Lawrence.
Haan W. de, 1833-1 850 - Crustacea. In: Fauna Japonica sive
Description Animalium, quae in Itinere per Japoniam,
Jussu et Auspiciis Superiorum, qui Summum in India
Botava Imperium Tenent, Suscepto, Annis 1823-1830
Collegit, Notis, Observationibus et Adumbrationibus
Illustravo. P. F. von Siebold & J. Miiller (eds). Lugduni-
Batavorum (=Leiden).
Huxley T. H., 1879 (1878) - On thè classification and thè
distribution of thè crayfishes. Proceedings of thè Scientific
Meetings of thè Zoological Society of London, London,
1878:752-788.
Latreille P. A., 1825 - Histoire Naturelle Entomologie ou
Histoire naturelle des Crustacés, des Arachnides et des
Insectes. In: Encyclopédie Méthodique. A. G. Desmarest
(ed.). Roret, Paris, 10: 1-344.
Lòrenthey E., 1902 - Neue Beitràge zur tertiàren Decapo-
denfauna Ungams. Mathematisch-Naturwissenschaftliche
Berichte aus Ungarn, Budapest, 18: 98-120.
Lòrenthey E. & Beurlen K., 1929 - Die fossilen Decapoden
der Lànder der Ungarischen Krone. Geologica Hungarica,
Serie Paleontologica, Budapest, 7 (3): 1-420.
Meyer H. von, 1 835 - Briefliche Mitteilungen. Neues Jahr-
buch fiir Mineralogie, Geologie, und Palaontologie,
Stuttgart.
Meyer H. von, 1842 - Uber die in dem dichten Jurakalk von
Aalen in Wiirtemburg vorkommenden Spezies des Crusta-
ceengenus Prosopon. Beitràge zur Petrefaktenkunde, Bay-
reuth, Heft 5: 70-75.
Meyer H. von, 1851 - Briefliche Mitteilgungen. Neues
Jahrbuch fiir Mineralogie, Geologie, und Palàontologie,
Stuttgart.
Meyer H. von, 1857 - Briefliche Mitteilungen. Jahrbuch
fiir Mineralogie, Geologie, und Palàontologie, Stuttgart.
Meyer H. von, 1860 - Die Prosoponiden oder die Familie der
Maskenkrebse. Palaeontographica, Stuttgart, 7: 183-222.
Mutiu R. & Bàdàlutà A., 1971 - La présence des décapodes
anomures et dromiacés dans les calcaires tithoniques
de Piate-Forme Moésienne. Annales Instituti Geolo¬
gici Publici Hungarici, Budapest, 54 (2): 245-525.
Ortmann A., 1892 - Die Abtheilungen Hippidea, Dromiidea,
und Oxystomata: Die Decapoden-Krebse des Strass-
burger Museums, mit besonderer Berucksichtigung der
von Herm Dr. Dòderlein bei Japan und bei den Liu-Kiu-
Inseln gesammelten und z.Z. im Strassburger Museum
aufbewahrten Formen. V. Theil. Zoologische Jahrbiicher,
Abtheilung fiir Systematik, Geographie, und Biologie der
Thiere, 6: 532-588.
Patrulius D., 1959 - Contributions à la systématique des déca¬
podes néojurassiques. Revue de Géologie et Géographie,
Bucharest, 3 (2): 249-257.
Patrulius D., 1966 - Les Décapodes du Tithonique inférieur
de Wozniki (Carpates Polonaises Occidentales). Annales
de la Société Géologique de Pologne, Kraków, 36 (4):
495-517.
Remes M., 1895 - Beitràge zur Kenntnis der Crustaceen
der Stramberger Schichten. Bulletin International de
l’Académie des Sciences de Bohème, Prague, 2: 200-204.
Reuss A. E., 1859 - Zur Kenntnis fossiler Krabben. Akademie
der Wissenschaften Wien, Vienna, Denkschrift 17: 1-90.
Samouelle G., 1819 - The Entomologist’s Usefìil Compendium,
or an Introduction to thè British Insects, etc. T. Boys, London.
Shirk A. M., 2006 - A novel assemblage of decapod Crus¬
tacea, ffom a Tithonian coral reef olistolith, Purcàreni,
Romania: systematical arrangement and biogeographical
perspective. Unpublished Master’s Thesis, Kent State
University, Kent, Ohio, USA.
Via Boada L., 1981 - Les Crustacés Décapodes du Ceno-
manien de Navarra (Espagne): Premiers résultats de
l’étude des Galatheidae. Géobios, Lyon, 14 (2): 247-251.
Carne E. Schweitzer - Dept. of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, U.S.A.
e-mail: cschweit@kent.edu
Rodney M. Feldmann - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rfeldman@kent.edu
Aubrey M. Shirk - Department of Geoscience, University of Nevada, 4505 Maryland Parkway, Las Vegas, Nevada 89154-4010, U.S.A.
e-mail: shirka2@unlv.nevada.edu
Iuliana Lazàr - Department of Geology and Palaeontology, 1, N. Balcescu Ave., sect. 1, 010041 Bucharest, Romania.
e-mail: iulia_lazar@geo.edu.ro
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Giorgio Teruzzi & Alessandro Garassino
Coleia Broderip, 1835 (Crustacea, Decapoda, Coleiidae)
from thè Mesozoic of Italy: an update
The discovery of Coleia Broderip, 1835, from thè
Lower Jurassic (Sinemurian) of Osteno (Lugano Lake,
N Italy) was reported for thè first time by Pinna (1968,
1969) who recorded nine specimens, referring them to C.
mediterranea Pinna, 1968, and C. viallii Pinna, 1968. The
first, well known by some very well preserved specimens,
shows thè following morphological characters: subrec-
tangular carapace with deep cervical and post-cervical
incisions; ocular incision with evident supra-antennal
spine; cervical groove deeper than branchiocardiac
groove; strong postcervical median carina and two strong
branchial carinae; pereiopod I longer than thè body length;
short triangular carpus; movable and fixed fingers with
distai extremity slightly curved; subtriangular telson with
median spine and two lateral spines at thè distai extrem¬
ity; petaloid uropodal exopod with rounded diaeresis. The
second, known by some complete specimens, shows thè
following morphological characters: subrectangular cara¬
pace; wide ocular incisions with supra-antennal spines
and separated by a shallow intraocular incision; narrow
cervical incisions and wide postcervical incisions; deep
cervical groove V-shaped; slender postrostral and pos-
torbital carinae; postcervical median carina stronger than
two branchial carinae; abdomen longer than carapace;
abdominal somites I-V with a strong and median tubercle;
chela of pereiopod I with movable finger bent at thè distai
extremity and as long as thè fixed fìnger; pereiopods II-IV
shorter than pereiopod I and with little chelae; triangular
telson with two tuberculate lateral carinae; petaloid uro¬
podal exopod with sinuous diaeresis.
Field researches in Osteno outcrop during thè eighties
and nineties of thè last century yielded many specimens
studied by Teruzzi (1990) who recorded 12 complete and
incomplete specimens, referring them to C. pianai Te¬
ruzzi, 1990, and C.popeyei Teruzzi, 1990.
The first shows thè following morphological char¬
acters: subrectangular carapace; almost straight frontal
margin with two little supraorbital spines; ocular inci¬
sions with supra-antennal spine; cervical incisions deeper
than postcervical incisions; cervical and branchiocardiac
grooves V-shaped; postcervical median carina stronger
than two branchial carinae; very elongate and thin pereio¬
pod I; chela of pereiopod I with movable finger at thè
distai extremity curved and longer than fixed finger; sub¬
triangular telson with distai extremity rounded and with
lateral carinae; and ovai uropodal exopod with rounded
diaeresis. The second species shows thè following mor¬
phological characters: subrounded carapace; almost
straight frontal margin with a wide intraocular incision;
cervical groove V-shaped; slender median postcervical
carina; two slender branchial carinae; pereiopod I with
strong and stout propodus; chela of pereiopod I with
movable and fixed fingers at thè distai extremity slightly
curved and similar in length; subtriangular telson with
two lateral carinae; and uropodal exopod with rounded
diaeresis.
Recently, Garassino & Gironi (2006) described 47
very well preserved specimens from Monte Verzegnis
(Udine) from thè Upper Triassic (Rhaetian), referring
them to Coleia boboi Garassino & Gironi, 2006, having
thè following morphological characters: subovoid cara¬
pace with two strong and elongate supraorbital spines;
two thin postorbitai carinae; thin median postcervical
carina; two thin branchial carinae; deep cervical groove;
deep cervical and postcervical incisions dividing thè
margin into three parts; chela of pereiopod I with mov¬
able and fixed fingers gently bent at thè distai extremity
and similar in length; subtriangular telson with two strong
tuberculate lateral carinae and spiny lateral margins; and
rounded uropodal exopod with rounded diaeresis.
Today, Coleia Broderip, 1835, known from thè Upper
Triassic (Camian) to thè Upper Jurassic (Tithonian) of
Europe (Germany, Great Britain, France, Russia and
Italy), Japan, and Madagascar, includes 22 species: 2
species from thè Upper Triassic: C. uzume Karasawa,
Takahashi, Doi & Ishida, 2003, and C. boboi Garassino,
& Gironi 2006; 18 species from thè Lower Jurassic: C.
antiqua Broderip, 1835; C. barrovensis (McCoy, 1849);
C. escheri (Oppel, 1862); C. edwardsi (Moriére, 1864); C.
wilmcotensis (Woodward, 1866); C. moorei (Woodward,
1866); C. brodiei (Woodward, 1866); C. crassichelis
(Woodward, 1866); C. morierei (Renault, 1889); C. tenui-
chelis Woods, 1925; C. bredonensis Woods, 1925; C.
sinuata Beurlen, 1928; C. sibirica Chemyshev, 1930; C.
theodorii Kuhn, 1952; C. viallii Pinna, 1968; C. mediter¬
ranea Pinna, 1968; C. pianai Teruzzi, 1990; C. popeyei
Teruzzi, 1990; and 2 species from Upper Jurassic: C. lon-
gipes (Fraas, 1855), and C. incerta Secretan, 1964.
References
Garassino A. & Gironi B., 2006 — Coleia boboi n. sp.
(Crustacea, Decapoda, Eryonoidea) from thè Late
Triassic (Rhaetian) of Monte Verzegnis (Udine, NE
Italy). Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale di Milano , Milano,
147(1): 93-102.
96
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Pinna G., 1968 - Gli Erionidei della nuova fauna
sinemuriana a crostacei decapodi di Osteno in
Lombardia. Atti della Società italiana di Scienze
naturali e del Museo civico di Storia naturale di
Milano, Milano, 107: 93-134.
Pinna G., 1969 - Due nuovi esemplari di Coleia viallii
Pinna, del Sinemuriano inferiore di Osteno in Lombar¬
dia (Crustacea, Decapoda). Annali del Museo di Storia
Naturale di Genova, Genova, 77: 626-632.
Teruzzi G., 1990 - The genus Coleia Broderip, 1835
(Crustacea, Decapoda) in thè Sinemurian of Osteno in
Lombardy. Atti della Società italiana di Scienze natu¬
rali e del Museo civico di Storia naturale di Milano,
Milano, 131 (4): 85-104.
Fig. 1 - A) C. boboi Garassino & Giorni, 2006. B) C. mediterranea Pinna, 1968. C) C. viallii Pinna, 1968. D) C. pinnai Teruzzi, 1990.
E) C. popeyei Teruzzi, 1990. (Drawings not in scale).
Giorgio Teruzzi - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: gteruz@tin.it
Alessandro Garassino - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: agarassino@libero.it
3rd Symposium on Mesozoic and Cenozoic Decapod Cnistaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
r
Francisco J. Vega, Fernando Alvarez & Gerardo Carbot-Chanona
Albian penaeoidea (Decapoda, Dendrobranchiata) from Chiapas,
southern Mexico
A new fossiliferous plattenkalk facies locality known
as E1 Chango has yielded numerous plant remains, fishes,
and crustaceans. E1 Chango is located approximately
30 km Southwest of Tuxtla Gutiérrez, Chiapas (Fig. 1).
Laminar dolomites of variable thickness (2 to 60 cm) crop
out in valleys of a karstic landscape. These deposits corre-
spond to thè base of thè Sierra Madre Formation, defined
as a 2,590 m sequence of Aptian to Santonian dolomites
and limestones (Fig. 2), deposited mainly on shallow
marine facies (Steele & Waite, 1986). Recently, another
age equivalent locality has been reported to contain plant
remains, insects, numerous and diverse crustaceans, as
well as fishes (Vega et al, 2006).
Fossils from E1 Chango include plant remains that
resemble Brachyphylluml sp., as well as other diverse
leaves (Figs. 3A-3D). Fishes of different species and
sizes are also found (Figs. 3E-3F), including macrosemiid
fishes and other groups with affinities with Asian and
south American forms (Ovalles-Damiàn et al, 2006).
Crustacean remains are rather scarce, and include pos-
sible dorippid crabs (Fig. 3G) and shrimps (Figs. 3H-3L),
whose preservation is usually very poor, but thè occur-
rence is significant because they represent thè first fossil
penaeoids described from Mexico.
C
.2
'S
o
Ph
<u
cd
ed
fc
<D
• W*
V
SZ3
■:%
A
2700 m
Rudist packestone s
_ 2200 m
Algal packestones
and wackestones
9
e
o
G
<u
U
<
G
G
CX
<
* A
r
r
Rudist wackestones
and packestones
_ 1500 m
c ° * 9 Q f
• &
e a
Coral-gastropod
wackestones
_700 m
Dolomite and
laminar dolomite
El Chango
.0 m
Fig. 1 - Location map of El Chango, Ocozocozutla County, Chiapas,
southern Mexico.
Fig. 2 - Stratigraphic section of thè Sierra Madre Formation, showing
relative position of deposits that crop out at El Chango.
Fig. 3 - A-B) Brachyphylluml sp., X 2.0. C-D) Unidentified plant remains, X 0.5. E-F) Unidentified fishes. E) X 0.3; F) X 5.0.
G) Dorippid? crab, hypotype IHNFG-2928, X 4.0. H-L) Unidentified crustaceans. H) Hypotype IHNFG-2929, X 5.0. 1) Lost specimen,
X 5.0 (scale in originai photograph). L) Flypotype IHNFG-2930, X 5.0.
3*° SYMPOSIUM ON MESOZOIC AND CENOZOIC DECAPOD CRUSTACEANS
99
This locality is similar in age and lithology to thè
one found near Profeti, Italy, from where palaeomo-
nids and carideans have been reported (Bravi & Gar-
assino, 2000).
Specimens are deposited in thè paleontological col-
lection of thè Museo de Paleontologia Eliseo Palacios
Aguilera, Instituto de Historia Naturai y Ecologia,
Tuxtla Gutiérrez, Chiapas, under acronym IHNFG.
Systematic paleontology
Superfamily Penaeoidea Rafinesque, 1815
Family Penaeidae? Rafinesque, 1815
Description. Rostrum short, probably reaching distai
portion of first antennular article. Dorsal surface of ros¬
trum with seven regularly spaced teeth, decreasing in size
anteriorly; posterior-most teeth above posterior margin of
orbit. Tip acute, simple. Ventral surface of rostrum smooth,
devoid of teeth. Carapace smooth, globose; dorsal surface
devoid of teeth beyond rostral ones. Antennal angle with
well defined spiniform projection. Pterygostomian angle
undefined. Ventral margin broadly rounded. Posterior
margin simple, straight. Abdomen smooth without obvi-
ous notches or grooves. First abdominal somite shortest,
0.6 times thè length of second somite; posterolateral
margin of pleuron overlapping second somite, ventral
margin rounded. Second somite approximately rectangu-
lar, posterior margin of pleuron slightly concave. Third
somite longer than first two, ventral margin rounded;
interlocking midlateral hinge present on posterolateral
margin of pleuron. Fourth somite of about same length as
third one, similar in shape; interlocking midlateral hinge
present on posterolateral margin of pleuron. Fifth somite
shorter than fourth, pleuron approximately rectangular;
interlocking midlateral hinge present on posterolateral
margin of pleuron. Sixth somite thè longest; dorsal,
ventral, and posterior margins of pleuron straight. Telson
shorter than uropods. Eyes well developed, cornea spheri-
cal. Antennule with peduncle shorter than scaphocerite,
antennular fìagellum long, about 4 times as long as total
length of organism. Scaphocerite 1.4 times as long as
antennular peduncle. Pereopods slender, ischia and carpi
of all five pairs very similar in width and length.
Material. One specimen, hypotype IHNFG-2931
(Fig.4).
Measurements. Carapace length = 41.7 mm, width =
11.1 mm.
Remarks. The present specimen is placed in thè
Penaeoidea based on its generai shape, a pleuron of thè
first abdominal somite that overlaps that of thè second
one, and five pereiopods of apparently thè same thick-
ness. Unlike modem species of penaeidae, thè rostral
teeth in thè Chiapas shrimp are all restricted to thè ante-
rior-most portion of thè rostrum anterior to thè posterior
margin of thè orbit. In modem penaeidae, there is always
an epigastric tooth, posterior to thè orbit, or a series of
teeth that reach thè posterior half of thè carapace along a
mid-dorsal carina (Pérez-Farfante & Kensley, 1997). We
compared our material with Jurassic and Cenomanian
penaeoideans from Solnhofen and Lebanon (Schweigert,
2001; Schweigert & Garassino, 2004; Garassino, 1994).
Those penaeoideans are different from thè Mexican
specimen in having rostra of different shape and length.
The Chiapas shrimp is similar in body proportions to thè
extant Farfantepenaeus and Litopenaeus, some of whose
species inhabit Mexican waters. The first abdominal
somite is short, thè rostrum lacks ventral teeth, unlike
modem penaeidae. Future field work may produce more
and best preserved specimens, in order to define if this
form represents a new genus.
Fig. 4 - Penaeidae? Flypotype IHNFG-2931, X 2.5. Photo and line drawing.
100
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
References
Bravi S. & Garassino A., 1 998 - Plattenkalk of thè Lower
Cretaceous (Albian) of Petina, in thè Albumi Mounts
(Campania, S Italy) and its decapod cmstacean assem-
blage. Atti della Società italiana di Scienze naturali e
del Museo civico di Storia naturale in Milano, 138:
89-118.
Bravi S. & Garassino A., 2000 - The Profeti platy
dolomite and its decapod cmstacean assemblage.
Studi e Ricerche, Associazione Amici del Museo -
Museo Civico “G. Zannato”, Montecchio Maggiore
(Vicenza), 2000: 11-13.
Garassino A., 1994 - The macruran decapod crustaceans
of thè Upper Cretaceous of Lebanon. Paleontologia
Lombarda, Nuova serie, Milano, III: 1-27.
Ovalles-Damiàn E., Alvarado-Ortega J. & Blaco-Pinón
A., 2006 - Los peces fósiles del Cretàcico Inferior de
Chiapas. In: X Congreso Nacional de Paleontologia,
México City, Memorias.
Pérez-Farfante I. & Kensley B., 1997 - Penaeoid and
Segerstoid Shrimps and Prawns of thè World. Keys
and Diagnoses for thè Families and Genera. Mémoires
du Museum national d'Histoire naturelle, 175: 1-233.
Rafinesque C. S., 1815 -Analyse de la Nature, ou Tableau
de FUnivers et des corps Organises. Palermo.
Schweigert G., 2001 - Eine neue Art der Gattung Antrim-
pos Miinster (Crustacea: Decapoda: Penaeidae) aus
dem Oberjura Siiddeutschlands. Stuttgarter Beitrage
zur Naturkunde, Series B, Stuttgart, 307: 1-33.
Schweigert G. & Garassino A., 2004 - New genera and
species of shrimps (Crustacea: Decapoda: Dendro-
branchiata: Caridea) from thè Upper Jurassic lithos-
tratigraphic limestones of S Germany. Stuttgarter Bei¬
trage fur Naturkunde, Serie B, Stuttgart, 350: 1-33.
Steele D. R. & Waite L. E., 1986 - Contributions to thè
stratigraphy of thè Sierra Madre Limestone (Creta¬
ceous) of Chiapas. México, Universidad Nacional
Autònoma, Instituto de Geologia, Boletin, 102: 1-175.
Vega F. J., Garcia-Barrera P., Perrilliat M. C., Coutino
M. A. & Marino-Pérez R., 2006 - E1 Espinal, a new
plattenkalk locality from thè Lower Cretaceous Sierra
Madre Formation, Chiapas, Southeastem Mexico.
Revista Mexicana de Ciencias Geológicas, 23 (3):
323-333.
Francisco J. Vega - Instituto de Geologia, UNAM, Ciudad Universitaria, Coyoacàn, México, D. F. 04510, Mexico.
e-mail: vegver@servidor.unam.mx
Fernando Àlvarez - Instituto de Biologia, UNAM, Ciudad Universitaria, Coyoacàn, México, D. F. 04510, Mexico.
e-mail: falvarez@servidor.unam.mx
Gerardo Carbot-Chanona - Museo de Paleontologia Eliseo Palacios Aguilera, Instituto de Historia Naturai y Ecologia,
Calzada de los Hombres Ilustres S/N, Parque Madero, Tuxtla Gutiérrez, Chiapas 29000, México.
e-mail: carbotsaurus@yahoo.com
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
James R. Verhoff, Pài M. Muller, Rodney M. Feldmann &
Carrie E. Schweitzer
A novel Paleocene decapod fauna from thè Kambiihel Formation
The Paleocene Kambiihel Formation of eastem Austria
preserves a decapod fauna that existed after thè K/T mass
extinction. The fauna contains a diverse assemblage of
brachyurans (true crabs) and anomurans (including squat
lobsters). These crabs and squat lobsters were reef-dwell-
ing organisms that were part of a community structure
that was preserved in rocks formed before and after thè
K/T extinction (Muller, 2004). This makes thè Kambiihel
Formation important to any attempt to understand thè
effect of thè K/T mass extinction on decapods.
Reefs were extremely favorable places for preserva-
tion of decapod crustaceans, specifically brachyurans and
anomurans (Muller, 2004). Several types of reefs have
been preserved in thè fossil record, with most of thè crab
material found in patch-reefs or coral-carpets (Muller,
2004). The Kambiihel Formation itself is probably a pre¬
served ffinging reef (Muller, 2004). Despite thè excellent
preservational potential, little is known about thè history
of decapods associated with reefs. This may be due to thè
small size of thè decapod fossils which makes them dif-
ficult to recognize, hard to collect, and of little interest to
collectors (Muller, 2004). The small size, combined with
thè relatively fragile carapace, of brachyurans makes them
difftcult to preserve, unless there is a sheltered area in
which thè carapaces can be embedded and buried (Muller,
2004). Decapod remains can also be swept off thè reef
and collected in fissures and cavities (Muller, 2004). Such
preservation does tend to create biases, however. Specifi¬
cally, thè remains are generally disarticulated, thè remains
are sorted by size, and more chelae than carapaces are
preserved (Muller, 2004). Also, it must be noted that, due
to thè patchy nature (both temporally and spatially) of thè
decapod reef faunas, it is impossible to identify whether
any change was graduai or punctuated (Muller, 2004).
To analyze decapod survival, it is important to look at
other taxa that were affected by thè K/T mass extinction.
The Kambiihel Formation has been interpreted as having
been deposited in a reef environment. The effects of thè
K/T extinction on corals, discussed below, were dramatic,
and thè impact of these changes on thè decapods found
in thè Kambiihel Formation is as yet unknown. Also, a
shift in plankton dominance is associated with thè K/T
mass extinction, which may have affected crabs and lob¬
sters indirectly. Finally, other survivors of thè K/T mass
extinction show important survival strategies with which
decapod survival can be compared.
Decapods as a taxon appear to have been remarkably
resistant to mass extinctions. Sixty-six percent of deca¬
pod genera in Denmark and Sweden survived thè K/T
mass extinction despite alterations of thè habitat (Col¬
lins & Jakobsen, 1994), and of thè 38 families studied
by Schweitzer & Feldmann (2005), 79% crossed thè K/T
boundary. Data on genera do show an extinction event,
but this event occurred over thè entire Late Cretaceous
(Schweitzer & Feldmann, 2005). This is not surprising, as
thè Cretaceous was a time of rapid radiation and extinc¬
tion for thè decapods (Schweitzer & Feldmann, 2005).
One study of Antarctic lobsters found that thè effects of
thè K/T mass extinction were either subtle or nonexistent
(Feldmann & Tshudy, 1989). Decapod faunas immedi-
ately following thè K/T mass extinction tended to be
relicts, rather than new or immigrant species (see Cope et
al., 2005; Feldmann & Tshudy, 1989, for examples). This
may be a result of thè Cretaceous radiation event. Large
numbers of newly evolved taxa in thè Late Cretaceous
coupled with relatively few extinctions at thè K/T bound¬
ary may have resulted in decapods occupying so many
niches that immigrant taxa could not form stable popula-
tions in new environments.
The traits found in decapods that survived thè K/T
event are similar to those found in survivors in other taxa.
Decapods that survived thè K/T extinction extended over
a broad geographic region and employed a generalist
feeding strategy (Schweitzer & Feldmann, 2005). These
behavioral traits are similar to those found in bivalve sur¬
vivors of thè K/T event. Deposit feeding bivalves, which
would have had a diet similar to generalists, survived better
than suspension feeding bivalves (Stilwell, 2003; Hansen
et al., 2004). Geographically widespread bivalves also
tended to survive better than those with more restricted
ranges (Stilwell, 2003). Decapods from temperate to high
latitudes also tended to have better survival rates, though
many taxa from subtropical environments and even from
near thè Chicxulub impact area in Yucutan, Mexico, sur¬
vived as well (Schweitzer & Feldmann, 2005). Similarly,
high-latitude bivalves had better survival rates than low-
latitude bivalves, although it is not clear whether bivalves
from high latitude regions survived because of latitudinal
effects or because of thè broad geographic ranges high
latitude bivalves occupied (Stilwell, 2003).
Bivalve survivors of thè K/T extinction event also
had several traits that form thè basis for comparison
with traits of decapod survivors. Infaunal bivalves seem
to have become more common after thè K/T extinction,
though thè cause is stili uncertain (see Lockwood, 2004
for further discussion). Omamentation on bivalves was
found to have no effect on survival during thè K/T extinc¬
tion (Kelley et al., 2001). However, bivalves use differ-
ent mechanisms than decapods to construct their shells.
Finally, thè traits that were favorable in bivalves during
thè recovery period were thè reverse of thè traits favored
in thè extinction-suspension feeders were favored in thè
recovery period, for example, while they were selected
against during thè extinction event itself (Stilwell, 2003).
102
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
The reefs of thè Kambuhel Formation were also
affected by thè K/T mass extinction. The K/T event had
two distinct effects on such coral reefs. The extinction
event selected against those corals that had forms similar
to modem reef-building corals that live in association
with zooxanthellae, with thè end result being extinction
of 30% +/- 8% of coral genera (Kiessling & Baron-Szabo,
2004). However, thè total volume of coral reef mass
actually increased in thè Paleocene, and corals became
more important as reef builders after thè K/T extinc¬
tion (Kiessling & Baron-Szabo, 2004). The Paleocene
increase in coral reef volume was concentrated in Europe,
north Africa, and south America (Kiessling & Baron-
Szabo, 2004). It is unknown what effect thè changes in
type and volume of coral had on decapods.
Plankton also played a role in thè formation of sub-
strate, and there was a high turnover in plankton at thè
K/T boundary. Specifically, there was a rise in biogenic
silica production associated with increased dominance of
diatoms and other silica-shelled plankton (Hollis, 2003).
Also, smaller species of plankton began to colonize lower
latitudes (Gartner, 1996). Both of these trends would
have affected thè substrate and food web of thè decapod
community in thè Kambuhel Formation, but thè specific
effects of thè plankton turnover on decapods are as yet
unknown.
In order to study thè impact of thè K/T extinction
and subsequent recovery on thè Kambuhel decapods,
thè decapods will be studied within thè framework of
extinction/recovery studies of other organisms. It is also
noted that decapods may show their own unique pattems
of recovery strategies, and thè Kambuhel decapods pro¬
vide an opportunity for such study.
References
Collins J. S. H. & Jakobosen S. L., 1994 - Synopsis of
thè biostratigraphic distribution of thè crab genera
(Crustacea, Decapoda) of thè Danian (Palaeocene) of
Denmark and Sweden. Bulletin of thè Mizunami Fossil
Museum, Mizunami, Japan, 21: 35-46.
Cope K. H., Utgaard J. E., Masters J. M. & Feldmann R.
M., 2005 - The fauna of thè Clayton Formation (Pa¬
leocene, Danian) of southern Illinois: A case of K/P
survivorship and Danian recovery. Bulletin of Mizu¬
nami Fossil Museum, Mizunami, Japan, 32: 97-108.
Feldmann R. M. & Tshudy D. M., 1989 - Evolution-
ary pattems in macrurous decapod cmstaceans from
Cretaceous to early Cenozoic rocks of thè James Ross
Island region, Antarctica. Geological Society Special
Publication, Boulder, Colorado, 47: 183-195.
Gartner S., 1996 - Calcareous nanofossils at thè Creta-
ceous-Tertiary Boundary, 27-47. In: Macleod N. &
Keller G. (eds.), Cretaceous-Tertiary Mass Extinc-
tions; Biotic and Environmental Changes. W W.
Norton and Company , New York, NY.
Hansen T. A., Kelley P. H. & Haasl D. M., 2004 - Pale-
oecological pattems in molluscan extinctions and
recoveries: comparison of thè Cretaceous-Paleogene
and Eocene-Oligocene extinctions in North America.
Palaeogeography, Palaeoclimatology, Palaeoecology,
Amsterdam, 214: 233-242.
Hollis C. J., 2003 - The Cretaceous/Tertiary boundary
event in New Zealand: profìling mass extinction. New
Zealand Journal of Geology and Geophysics, New
Zealand, 46: 307-321.
Kelley P. H., Hansen T. A., Graham S. E. & Huntoon
A. G., 2001 - Temporal pattems in thè efficiency of
naticid gastropod predators during thè Cretaceous
and Cenozoic of thè United States Coastal Plain. Pal¬
aeogeography, Palaeoclimatology, Palaeoecology,
Amsterdam, 166: 165-176.
Kiessling W. & Baron-Szabo R. C., 2004 - Extinction
and recovery pattems of scleractinian corals at thè
Cretaceous-Tertiary boundary. Palaeogeography,
Palaeoclimatology, Palaeoecology, Amsterdam, 214:
195-223.
Lockwood R., 2004 - The K/T event and infaunality;
morphological and ecological pattems of extinction
and recovery in veneroid bivalves. Paleobiology,
Lawrence, Kansas, 30 (4): 507-521.
Muller P. M., 2004 - History of reef-dwelling decapod
cmstaceans from thè Palaeocene to thè Miocene with
comments about Mesozoic occurrences. Fòldtani
Kòzlòny, Budapest, 134 (2): 237-255.
Schweitzer C. E. & Feldmann R. M., 2005 - Decapod
cmstaceans, thè K/P event, and Palaeocene recovery,
17-53. In: Koenemann S. & Jenner R.A. (eds.), Crus¬
tacea and Arthropod Relationships. Taylor & Francis
Group , New York, NY.
Stilwell J. D., 2003 - Pattems of biodiversity and faunal
rebound following thè K-T boundary extinction event
in Austral Palaeocene molluscan faunas. Palaeogeog¬
raphy, Palaeoclimatology, Palaeoecology, Amster¬
dam, 195: 319-356.
James R. Verhoff - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: jverhoff@kent.edu
Pai M. Muller - Magyar Aliami Fòldtani Intézet, Stafània ut 14, 1143, Budapest, Hungary.
e-mail: muller.pal@freemail.hu
Rodney M. Feldmann - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rfeldman@kent.edu
Carrie E. Schweitzer - Dept. of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, U.S.A.
e-mail: cschweit@kent.edu
3rd Symposium on Mesozoic and Cenozoic Decapod Cmstaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
David A. Waugh, Rodney M. Feldmann, Angie Hull, Kristie Hein &
Carrie E. Schweitzer
Scaling and classifìcation of cuticular pits in raninids
The range of morphology, size, and density of pits
observed on thè surface of raninid crabs is examined
with thè goal of creating biologically meaningful
characters that can be coded for future phylogenetic
analysis. In addition to morphology of thè pits, thè size
and density of these pits is an obvious starting place in
thè search for characters; therefore, emphasis is placed
on understanding thè growth of these pits in relation to
carapace size.
Cuticle samples from extant and fossil raninids have
been obtained during an ongoing study of thè role that
cuticle microstructures may play in providing characters
when constructing decapod phylogenies. Material was
examined directly or cast if thè material was too rare
for destructive sampling. Extant cuticle from museum
spirit collections was cast or a section was extracted
from thè branchial region of thè carapace. Casts and
originals were examined using thè SEM or were whit-
ened with ammonium chloride and photographed with
a light microscope. Sampling of thè branchial region of
thè dorsal carapace allowed direct comparison across
taxa.
Morphologically, pits on thè surface of thè raninids
that were examined can be classified into six groups:
simple closed pits, closed pits with defined sides, closed
pits with small undefined nodes, perforations, pits with
setae, and pits with developed nodes and setae (Fig. 1).
The six groups have been Consolidated into five proposed
characters by combining Perforations and Pits with Setae.
Subsequent work will apply these characters and others
under development to a phylogenetic analysis of thè
Raninidae.
Twenty species in eight raninid genera have been
examined to determine pit size and density. In most thè
raninids observed to date, thè size of thè pits is observed
to increase, and thè density of pits to decrease, as thè car¬
apace becomes larger. The clear outliers in this trend are
thè samples of thè Cretaceous Eumorphocorystes sculp-
tus Binkhorst, 1857 and Notopocorystes stokesi (Mantell,
1 844) that show a significanti increased density of pits
in relation to their carapace size.
The link between thè expanding size and decreasing
density of pits (or nodes) can be expected from a simple
model in which a portion of cuticle starts out with a given
number and size of features, thè size of which expands
and density decreases with each molt. This model of
growth is too simple to apply blindly, and thè introduction
of smaller nodes between larger ones on thè surface of
Ramina ramina (Linnaeus, 1758) clearly does not follow
it. Future work will examine thè allometric relationship
of all cuticle features with thè expectation that certain
features will show no growth, others will expand with
growth, and some new features will be introduced during
growth.
Thus, it is possible to conclude that five proposed
character States can be described for coding raninid pits
in phylogenetic analysis. The size of pits on thè surface
of raninids tends to increase with thè growth of thè cara¬
pace; a corresponding decrease in thè density of thè pits is
also observed. Understanding these growth trends allows
classifying structures as large or small based upon aver-
age raninid growth trends. Understanding this scaling
also prevents features that are thè same relative size, but
present on carapaces of differing sizes, from being used
as evidence of taxonomic dissimilarity. Pit size appears
to be highly correlated with thè amount of carapace
growth; this trend can be marked on thè carapace of a
single specimen. Perforations observed in Eumorphoco¬
rystes sculptus and Notopocorystes stokesi exhibit much
higher densities than closed pits observed in thè bulk of
thè raninids. Unlike thè pits without setae in thè raninids,
setal pits observed in Callinectes sapidus Rathbun, 1 896,
a portunid crab used for comparison, do not grow as thè
carapace expands during ontogeny. Observing growth of
cuticle surface features provides a biologically meaning¬
ful way of separating classes of character States.
References
Adams A. & White A., 1848 - Crustacea. In: The Zool-
ogy of thè Voyage of H. M. S. Samarang, 1843-1846.
London. Adams, A. (ed.).
Binkhorst J. T. van, 1857 - Neue Krebse aus der Maes-
trichter Tuffkreide. Verh. Naturhist. Ver preuss.
Rheiml. Westf 14: 107-110.
Haan W. de, 1833-1850 - Crustacea. In: Fauna Japonica
sive Description Animalium, quae in Itinere per Japo-
niam, Jussu et Auspiciis Superiorum, qui Summum in
India Botava lmperium Tenent, Suscepto, Annis 1823-
1830 Collegit, Notis, Observationibus et Adumbratio-
nibus Illustravit. P. F. von Siebold & J. Miiller (eds.).
Lugduni-Batavorum (=Leiden).
Linnaeus C., 1758 - Systema Naturae per Regna Trio
Naturae, Secundum Classes, Ordines, Genera, Spe¬
cies, cum Characteribus, Differentiis, Synonymis,
Locis (edit. 10). Voi. 1. Holmiae (Stockholm), Lau-
rentii Salvii.
Mantell G. A., 1844 - The Medals of Creation, Or, First
Lessons in Geology, and in thè Study of Organic
104
EDITED BY ALESSANDRO GARASSINO, RODNEY M. FELDMANN & GIORGIO TERUZZI
Remains. Henry G. Bohn, York Street, Covent Garden,
London. 2: 457-1016.
Rathbun M. J., 1896 - The genus Callinectes. Proceed-
ings of thè United States National Museum, 1 8 ( 1 070):
349-375.
Rathbun M. J., 1933 - Descriptions of new species of
crabs from thè Gulf of California. Proceedings of thè
Biologica l Society of Washington , Washington, 46:
147-150.
Sakai T., 1963 - Description of two new genera and 14
new species of Japanese crabs from thè collection
of His Majesty thè Emperor of Japan. Crustaceana,
Leiden, 5 (3): 213-233.
Fig. 1 - Pits observed on thè surface of raninids. A) Simple closed pits, Lyreidus trìdentatus De Haan, 1841, extant. B) Closed pits with
defined sides, observed in Macroacaena sp., Eocene-lower Oligocene. C) Closed pits with undefined nodes, Raninoides louisianensis
Rathbun, 1933, extant. D) Cast of setal hairs in Umalia orientalis (Sakai, 1963), extant, poor definition of seta due to limitations of
casting technique and high magnification. E) Perforations, Eumorphocorystes sculptus Binkhorst, 1857, Cretaceous. F) Pits with setae
and nodes, Cosmonotus grayii Adams and White, 1 848, extant.
David A. Waugh - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: dwaugh@kent.edu
Rodney M. Feldmann - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: rfeldman@kent.edu
Angie Hull - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: acollmar@kent.edu
Kristie Hein - Department of Geology, Kent State University, Kent, Ohio 44242, U.S.A.
e-mail: khein@kent.edu
Carrie E. Schweitzer - Dept. of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, U.S.A.
e-mail: cschweit@kent.edu
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans - Museo di Storia Naturale di Milano, May 23-25, 2007
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo II
Ili - PELOSIO G., 1 968 - Ammoniti del Lias superiore (Toarciano) dell’Al¬
pe Turati (Erba, Como). Generi Hildoceras, Phvmatoceras, Paronice-
ras e Frechiella. Conclusioni generali, pp. 143-204, 2 figg., 6 taw.
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 taw.
II - MONTANARI L., 1969 - Aspetti geologici del Lias di Gozzano (Lago
d’Orta )./?/?. 23-92, 42 figg., 4 taw. n.t.
Ili - PETRUCCI F., BORTOLAMI G. C. & DAL PIAZ G. V., 1970 - Ri¬
cerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul
suo substrato cristallino, pp. 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, pp. 5-46, 2 tabb. n.t.
II - PINNA G. & LEVI-SETTI F., 1971 - I Dactylioceratidae della Provin¬
cia Mediterranea (Cephalopoda Ammonoidea). pp. 47-136, 21 figg.,
12 taw.
Ili - PELOSIO G., 1973 - Le ammoniti del Trias medio di Asklepieion (Ar-
golide, Grecia) - I. Fauna del «calcare a Ptychites » (Anisico sup.). pp.
137-168, 3 figg., 9 taw.
I
Volume XX
I - CORNAGGIA CASTIGLIONI O., 1971 - La cultura di Remedello.
Problematica ed ergologia di una facies dell’Eneolitico Padano, pp.
5-80, 2 figg., 20 taw.
II - PETRUCCI F., 1972 - Il bacino del Torrente Cinghio (Prov. Parma).
Studio sulla stabilità dei versanti e conservazione del suolo, pp. 81-127,
37 figg., 6 carte tematiche.
III - CERETTI E. & POLUZZI A., 1973 - Briozoi della biocalcarenite del
Fosso di S. Spirito (Chieti, Abruzzi). pp. 129-169, 18 figg., 2 taw.
Volume XXI
I - PINNA G., 1 974 - 1 crostacei della fauna triassica di Cene in Val Seriana
(Bergamo), pp. 5-34, 16 figg., 16 taw.
II - POLUZZI A., 1975 - 1 Briozoi Cheilostomi del Pliocene della Val d’ Ar¬
da (Piacenza, Italia). pp. 35-78, 6 figg., 5 taw.
III - 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 taw.
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 taw.
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 taw.
II - POLUZZI A., 1982 - 1 Radiolari quaternari di un ambiente idrotermale
del Mar Tirreno, pp. 45-72, 3 figg., 1 lab., 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., 1984 - Osteologia di Drepanosaurus ungule audatus, 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.
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,
575 figg.
Volume XXVI
I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia, Placo-
dontia).pp. 1-21, 23 figg.
II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara preisto¬
rico: dati e interpretazioni, pp. 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 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., 1999 - L’arte rupestre dell’Eritrea. Repertorio ragionato
ed esegesi iconografica, pp. 1-174, 268 figg.
Volume XXX
I - PEZZOTTA F. edited by, 2000 - Mineralogy and petrology of shallow
depth pegmatites. Paper ffom 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, pp. 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). pp. 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.
Ili - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphargis dalsassoi n. gen.
n. sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano
(Lombardy, N Italy). pp. 1-33, 19 figs., 1 tab., 3 pls.
Volume XXXII
I - ALESSANDRELLO A., BRACCHI G. & RIOU B„ 2004 - Polychaete,
sipunculan and enteropneust worms from thè Lower Callo vian (Middle
Jurassic) of La Voulte-sur-Rhòne (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.
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). 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. 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. Part 1. Infraorders Astacidea, Thalassinidea and Palinura.
pp. 1-64, 12 figs., 20 pls.
II - FUCHS D., 2006 - Morphology, taxonomy and diversity of vampyropod
Coleoids (Cephalopoda) from thè Upper Cretaceous of Lebanon. pp.
1-28, 9 figs., 9 pls.
III - CALDWELL M. W., 2006 - A new species of Pontosaunis (Squamata,
Pythonomorpha) from thè Upper Cretaceous of Lebanon and a phylo-
genetic analysis of Pythonomorpha. pp. 1-42, 18 figs., 1 pi.
Volume XXXV
I - DE ANGELI A. & GARASSINO A., 2006 - Catalog and bibiiography of
thè fossil Stomatopoda and Decapoda from Italy. pp. 1-95.
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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?oo<SL
MEMORIE
della Società Italiana
di Scienze Naturali
e del Museo Civico
Volume XXXV - Fascicolo III di Storia Naturale di Milano
MCZ
LIBRARY
STEFANIA NOSOTTI JAN 0 2 ZOOb
HARVARD
UNIVERSITY
TANYSTROPHEUS LONGOBARDICUS
(REPTILIA, PROTOROS AURI A) :
RE-INTERPRETATION S OF THE ANATOMY BASED ON NEW
SPECIMENS FROM THE MIDDLE TRIASSIC OF BESANO
(LOMBARDY, NORTHERN ITALY)
MILANO NOVEMBRE 2007
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
Felix: Felis jacobita (Com.). 9 pp., I tav.
II - MAGNI-GRIFFI F., 1865 - Di una specie d'Hippolais nuova per l’Ita¬
lia. 6 pp., 1 tav.
III - 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., 1 865 - 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 L, 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 - 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 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., 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 taw.
Volume III
I - EMERY C., 1873 - Studii anatomici sulla Vipera Redii. 16 pp., 1 tav.
II - GARO VAGLIO S., 1867 - Thelopsis, Belonia, Weitenweber a 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 - ÓARO VAGLIO S., 1871 - De Pertusariis Europae mediae commenta¬
no. 40 pp., 4 taw.
Volume IV
I - D’ ACHIARDI A., 1 868 - Corallarj fossili del terreno nummulitico del-
l’Alpi venete. Parte 1 1 . 32 pp. 8 taw.
II - GARO VAGLIO S., 1868 - Octona Lichenum genera vel adhuc con¬
troversa, vel 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 - MARTORELLI G., 1898 - Le forme e le simmetrie delle macchie nel
piumaggio. Memoria ornitologica. 112 pp., 63 figg., 1 taw.
Ili - PAVESI P„ 1901- L’abbate Spallanzani a Pavia. 68 pp., 14 figg., I tav.
Volume VII
I - DE ALESSANDRI G., 1910 - Studi sui pesci triasici della Lombardia.
164 pp., 9 tavv.
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 - AIRAGHI C., 1917 - Sui molari d’elefante delle alluvioni lombarde,
con osservazioni sulla filogenia e scomparsa di alcuni Proboscidati. pp.
187-242, 4 figg., 3 taw.
Volume IX
I - BEZZI M., 1918 - Studi sulla ditterofauna nivale delle Alpi italiane, pp.
1-164, 7 figg. 2 taw.
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.
III - 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)./?/?. 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./?/?. 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 - 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), 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.
Ili - VIALLI V., 1959 - Ammoniti sinemuriane del Monte Albenza (Berga¬
mo)./?/?. 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./?/?. 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.
III - ZANZUCCHI G., 1963 - Le Ammoniti del Lias superiore (Toarciano)
di Entratico in Val Cavallina (Bergamasco orientale), pp. 99-146, 2
figg.. 8 taw.
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
dintorni di Orosei (Sardegna)./?/?. 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, 27 figg. 9 taw.
II - DIENI I. e MASSARI F„ 1966 - Il Neogene e il Quaternario dei dintorni
di Orosei (Sardegna)./?/?. 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 taw.
Volume XVI
I - CARETTO P. G„ 1967 - Studio morfologico con l’ausilio del metodo
statistico e nuova classificazione dei Gasteropodi pliocenici attribuibili
al Mure.x brandaris Linneo./?/?. 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 tavv.
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)./?/?. 71-142.
5 figg., 11 taw.
Stefania Nosotti
Sezione di Paleontologia, Museo Civico di Storia Naturale di Milano
M CZ
L/SR/^pfy
JAN 02 2008
Tanystropheus longobardicus (Reptilia, Protorosauria):
re-interpretations of thè anatomy based on new specimens
from thè Middle Triassic of Besano (Lombardy, northern Italy)
1857 - 2007
150 ANNI
DI NATURA
E SCIENZA
SOCIETÀ ITALIANA
DI SCIENZE NATURALI
Volume XXXV - Fascicolo III
Novembre 2007
Memorie della Società Italiana di Scienze Naturali
e del Museo Civico di Storia Naturale di Milano
INDEX
INTRODUCTION . Pag. 4
INSTITUTIONAL ABBREVI ATIONS . Pag. 6
ANATOMICAL ABBREVIATIONS . Pag. 6
MATERIALS . Pag. 7
METHODS . Pag. 8
DESCRIPTION OF THE SPECIMENS . Pag. 9
Specimen MSNM BES SC 265 . Pag. 9
Skull . Pag. 10
Axial skeleton . Pag. 13
Appendicular skeleton . Pag. 1 5
Specimen MSNM BES SC 1018 . Pag. 20
Skull . Pag. 20
Axial skeleton . Pag. 27
Appendicular skeleton . Pag. 29
Specimen MSNM V 3663 a b . Pag. 38
Specimen MSNM BES 215 . Pag. 38
Specimen MSNM BES 351 . Pag. 41
Specimen MSNM V 3730 . Pag. 42
DISCUSSION . Pag. 44
Discussion and interpretation of thè skull anat-
omy of Tanystropheus : a new reconstruction ... Pag. 44
The axial skeleton: an overall description . Pag. 62
Cervical vertebrae and ribs . Pag. 62
Dorsal vertebrae and ribs . Pag. 67
Sacrai vertebrae . Pag. 69
Caudal vertebrae . Pag. 69
The mobility of thè vertebral column in Tanys¬
tropheus . Pag. 70
Gastralia . Pag. 71
Could Tanystropheus walk? . Pag. 71
The pes in Tanystropheus . Pag. 7 1
The hindlimb: terrestrial versus aquatic locomo-
tion . Pag. 73
Tanystropheus mode of life: on what side of thè
shoreline? . Pag. 76
IMPLICATIONS FOR PHYLOGENETIC
ANALYSIS . Pag. 80
CONCLUSIONS . Pag. 82
AKNOWLEDGEMENTS . . . Pag. 84
REFERENCES . Pag. 85
© 2007 Società Italiana di Scienze Naturali
Museo Civico di Storia Naturale di Milano
Corso Venezia ,55 -20121 Milano
In copertina: Tanystropheus longobardicus, MSNM BES SC 1018, skull. Watercolor by Massimo Demma.
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 - Novembre 2007 ISSN 0376-2726
A
Stefania Nosotti
Tanystropheus longobardicus (Reptilia, Protorosauria):
re-interpretations of thè anatomy based on new specimens
from thè Middle Triassic of Besano (Lombardy, northern Italy)
This monograph is dedicateci
to Giorgio Teruzzi and Anna Alessandrello,
far their unconditional trust and support
to Massimo Demmo,
whose generosity, patience, dedication and extreme professionalism
made an important contribution to this publication
to Dr. Rupert Wild,
who established a milestone in Tanystropheus research
Abstract - After more than one century since thè first report of Tanystropheus longobardicus from thè Middle Triassic of Besano,
new specimens from thè same outcrops are described. These specimens include two articulated skeletons and an isolated pes, all from
small-sized individuals, and fragmentary remains of larger individuals, i.e. a skull with some associated cervical vertebrae, an isolated
dorsal vertebra and isolated cervical ribs.
This new material confirms thè presence of T. longobardicus in thè Besano Formation, which previously yielded few evidence.
Moreover, it includes remarkably complete and well preserved specimens which provided thè opportunity of a new interpretation of
thè anatomy of Tanystropheus , formerly described on thè basis of a rich sample from thè Swiss Grenzbitumenzone.
The description presented here applies to small-sized individuals of Tanystropheus, traditionally interpreted as thè juveniles of
T. longobardicus. However, thè point is raised that they might represent thè adults of a different species, demonstrating thè presence
of two taxa among thè Swiss and Italian material referred to T. longobardicus. The holotype, and thè single known specimen, of thè
small-sized Tanystropheus meridensis from thè Meride Limestone is also considered and re-interpreted, leading to thè conclusion that
this species is probably a junior synonym of T. longobardicus.
Comparisons of thè specimens of Tanystropheus from thè Besano Formation with those from thè equivalent Grenzbitumenzone
helped to find thè problematic elements of thè classical reconstruction.
A new reconstruction of thè skull of Tanystropheus is presented based on a three-dimensional clay model, with a re-interpretation
of thè pre-orbital region, thè skull roof, and thè lower jaw. The reconstruction of thè temporal region of thè skull is shown to be highly
problematical. Finally, thè new specimens confimi thè presence of a sclerotic ring in Tanystropheus.
In thè postcranial skeleton, thè more important new information concem thè morphology of thè appendicular skeleton, which is
remarkably well preserved in thè new specimens. In particular, preservation of complete and perfectly articulated manus and pedes for
thè first time yields unequivocal evidence on their morphology.
The anatomy of thè appendicular skeleton, in particular that of thè hindlimb, is discussed in thè context of locomotion mode. An
overall view of previous studies on thè mode of fife of Tanystropheus is presented and discussed. According to these results, Tany¬
stropheus should be regarded as a marine protorosaur, with dose terrestrial ancestors, living in shallow waters. The feeding strategy of
Tanystropheus is discussed, on thè assumption that it likely was a slow, axial or paraxial swimmer with a stiff neck.
In conclusion, thè new information obtained from thè specimens described here is evaluated in thè context of thè recent cladistic
analyses of protorosaurian relationships, highlighting thè hearing of systematic anatomical work of originai materials on thè descrip¬
tion and coding of phylogenetically informative characters.
Key words: Tanystropheus, Tanystropheus longobardicus. Middle Triassic, Besano, northern Italy, new specimens, Tanystropheus
meridensis, anatomy, skull, pes, mode of life, systematic.
Riassunto - Tanystropheus longobardicus (Reptilia, Protorosauria): re-interpretazioni dell’anatomia basate su nuovi esemplari
provenienti dal Triassico medio di Besano (Lombardia, Italia settentrionale).
A distanza di oltre un secolo dalla prima segnalazione di Tanystropheus longobardicus negli strati fossiliferi del Triassico medio
di Besano (Varese), vengono qui descritti nuovi esemplari provenienti dai medesimi affioramenti. Si tratta di due scheletri in larga
parte articolati e di un piede isolato ascrivibili ad individui di piccola taglia e di resti frammentari di individui di grandi dimensioni, in
particolare un cranio associato ad alcune vertebre cervicali, una vertebra dorsale isolata e frammenti di coste cervicali.
Questo consistente nucleo di esemplari costituisce una ricca documentazione della presenza di T. longobardicus negli strati fos¬
siliferi di Besano. Tale presenza era precedentemente testimoniata solamente dall’olotipo, distrutto nel corso della Seconda Guerra
Mondiale, e da due esemplari frammentari conservati nelle Collezioni del Palaontologischen Institut und Museum der Universitàt,
Zurich (PIMUZ). La completezza e l’ottimo stato di conservazione dei nuovi materiali consente una re-interpretazione dell’anatomia
scheletrica di Tanystropheus, descritta nel secolo scorso sulla base di svariati esemplari affiorati sul versante svizzero del giacimento.
4
STEFANIA NOSOTTI
La nuova descrizione dell’anatomia di T. longobardicus qui presentata è applicabile ad individui di piccola taglia, che non supe¬
rano i due metri di lunghezza complessiva. Nell’interpretazione classica, tali esemplari rappresenterebbero gli stadi giovanili della
specie. Viene tuttavia avanzata l’ipotesi che in alternativa essi rappresentino gli adulti di una specie differente e che nel materiale
italiano e svizzero proveniente dalla Formazione di Besano siano in realtà presenti due specie, Luna di piccola taglia, l’altra di taglia
medio-grande. Viene anche preliminarmente re-interpretato l’olotipo, ed unico esemplare conosciuto, di Tanystropheus meridensis,
proveniente dal Calcare di Meride, concludendo che esso è probabilmente sinonimo di T. longobardicus.
11 dettagliato confronto dei nuovi esemplari provenienti dalla Formazione di Besano con quelli dell’equivalente Grenzbitumenzone
conservati nelle Collezioni del PIMUZ ha permesso di ottenere un’informazione più completa e di individuare gli aspetti problematici
della ricostruzione classica.
Viene presentata una nuova ricostruzione del cranio di Tanystropheus ottenuta tramite la realizzazione di un modello tridimen¬
sionale. La regione pre-orbitale del cranio, il tetto cranico e la mandibola sono stati re-interpretati con risultati innovativi e ragione¬
volmente attendibili, mentre la ricostruzione della regione temporale, rivelatasi particolarmente problematica, resta ipotetica e viene
pertanto proposta come un punto di partenza per ulteriori discussioni. I nuovi esemplari hanno tra l’altro definitivamente confermato
la presenza di un anello sclerotico nell’occhio di Tanystropheus.
Circa lo scheletro postcraniale, gli esemplari di Besano hanno fornito nuove rilevanti informazioni. In particolare, l’eccezionale
conservazione degli arti, unica nell’ambito di tutto il materiale conosciuto, ha permesso per la prima volta di accertare inequivocabil¬
mente alcune delle loro caratteristiche anatomiche.
L’anatomia dello scheletro appendicolare, soprattutto del piede, viene interpretata in relazione alla locomozione terrestre ed
acquatica. Questo, ed altri aspetti cruciali per l’interpretazione del modo di vita di Tanystropheus, vengono presentati e discussi, con
un excursus che include anche gli studi più recenti e il confronto con altri vertebrati estinti ed attuali. Nell’interpretazione dell’autore
Tanystropheus era un rettile marino nell’intero arco di vita, tuttavia non altamente specializzato per la vita acquatica in quanto stret¬
tamente imparentato con antenati francamente terrestri. Poiché Tanystropheus viene considerato un nuotatore lento dotato di un collo
piuttosto rigido, uno degli aspetti più problematici nell’interpretazione del suo modo di vita rimane quello della strategia utilizzata
per la cattura di prede presumibilmente molto sfuggenti. Contenuti stomacali a pesci e cefalopodi sono stati infatti rinvenuti in alcuni
esemplari nelle Collezioni del PIMUZ.
Infine l’informazione inedita ottenuta con lo studio dei nuovi esemplari viene valutata nel contesto delle più recenti analisi cla-
distiche concernenti le relazioni filogenetiche dei protorosauri e si dimostra come in alcuni casi la descrizione e la codificazione dei
caratteri siano basate su illazioni speculative presentate come dati certi, risultando di conseguenza errate e fuorvianti. Viene pertanto
sottolineata l’importanza dello studio anatomico di materiale originale per una descrizione e codificazione accurata dei caratteri filo¬
geneticamente informativi.
Parole chiave: Tanystropheus, Tanystropheus longobardicus. Triassico medio, Besano, Italia settentrionale, nuovi esemplari,
Tanystropheus meridensis, anatomia, cranio, pes, modo di vita, sistematica.
INTRODUCTION
The genus Tanystropheus was erected by Hermann
von Meyer (1847-1855) in 1852 (Quenstedt, 1963; Wild,
1976; I.C.Z.N., 1981), to comprise thè species T. conspic-
uus, based on isolated bones from thè Upper Muschelkalk
of Bayreuth. These elements were interpreted by von
Meyer as strikingly elongate caudal vertebrae of a reptile.
Von Meyer did not explain thè ethymology of thè name
Tanystropheus, which means “long ribbon”, derived from
thè Greek “tany-”= prefìx, meaning “long”, “stretched”
and “stróphos”= “strap”, “rope”, “ribbon”, referring to
thè slender, elongate shape of thè bones. According to
von Meyer, thè same bones had earlier been interpreted
by Georg zu Miinster as belonging to thè limb of a new
reptile, for which he had proposed thè name Macroscelo-
saurus (with no indication of thè relevant specific epithet,
if any). However, von Meyer gave no reference to any
publication by Miinster. He introduced thè name Tanys¬
tropheus maintaining that thè name given by Miinster was
no longer accepted, with no further explanation. Since
then, thè name Tanystropheus has almost exclusively
been used (Wild, 1976: 124), while only few authors
(Broili, 1915: 51; Kuhn, 1934: 1 1 8) considered thè use of
thè name Tanystropheus against thè principle of priority,
thè senior synonym being Macroscelosaurus .
Oskar Kuhn (1934) referred to “ Macroscelosaurus
Miinster, 1834” but no mention of Macroscelosaurus has
ever been found either in thè paper of 1834, or in any
other known published paper by Miinster (Kuhn, 1963:
5; Wild, 1974: 147; 1976: 124). Possibly, as suspected by
Wild R. (pers. comm., 1998), Miinster proposed thè name
in an unpublished private letter to von Meyer. Seemingly,
von Meyer first published Macroscelosaurus, as a junior
synonym of Tanystropheus. Following an application by
Wild (1975; 1976), thè International Commission on Zoo-
logical Nomenclature (I.C.Z.N., 1981) under its plenary
powers conserved Tanystropheus von Meyer, [1852] as
thè valid name for thè genus and suppressed Macroscelo¬
saurus von Meyer, [1852]. Thus von Meyer was accepted
as thè author, and 1952 as thè publication date of both
names.
In 1 886 Francesco Bassani reported a new reptile from
thè Middle Triassic of Besano (Varese Province, Lom-
bardy, Italy), and named it Tribelesodon longobardicus,
in reference to its tricuspid dentition. The specimen,
housed in thè Collections of thè Museo di Storia Naturale
di Milano, consisted of incomplete remains of a skull
and some associated postcranial elements, preserved on
a part and counterpart piate (Nopcsa v., 1923: 161, pi.
II). Bassani gave a very short preliminary description
of thè specimen, which at thè time was very poorly pre-
pared (Peyer, 1931: 83), and tentatively concluded that it
represented a flying reptile. As he did not include in his
description any illustration, it is difficult to understand
thè reasons of his interpretation. Bassani also referred to a
“second specimen” but Peyer (1931: 94) pointed out that
a second Tribelesodon specimen could not be located in
Milano, and conjectured that thè “second specimen” was
in fact a poorly preserved specimen of Macrocnemus.
Bassani never published thè monographic description of 1
Tribelesodon, and for more than fifteen years thè reptile
from Besano lay almost forgotten in thè Collections of thè
Museo di Storia Naturale di Milano.
The first detailed study of thè holotype of Tribelesodon
longobardicus was undertaken by Franz von Nopcsa in
A
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
5
1902 and 1904, and supported by grants from thè Wiener
Akademie der Wissenschaften (Nopcsa v., 1923: 161).
The results of thè study, and thè photographs of thè speci¬
men were not published until 1923.
In thè meantime, Gustav von Arthaber (1921) pub¬
lished a short description and a preliminary reconstruc-
tion of thè skull of Tribelesodon. Von Nopcsa provided
von Arthaber with a photograph of thè skull, that was also
published. At that time Tribelesodon was stili consid-
ered a flying reptile. Later, von Arthaber (1922: fig. 3 a)
published a composite photograph of thè whole speci¬
men, obtained by overlapping thè negatives of thè part
and counterpart taken by von Nopcsa in 1902 (Nopcsa v.,
1923: 161), and thè reconstruction of thè skull alreadly
published in 1921.
Von Nopcsa (1 923) continued to consider Tribelesodon
as a flying reptile (Nopcsa v., 1923: fig. 6). His reasoning
based on thè interpretation of some problematic, very
elongate bones as segments of a forelimb with a long digit
that he presumed supported a wing membrane. Once this
assumption had been made, it influenced von Nopcsa’s
interpretation of many of thè other preserved elements.
Even so, von Nopcsa did consider thè possibility that thè
problematic bones could be elongate vertebrae, similar to
thè cervical vertebrae of thè pterosaur Doratorhyncus , or
to thè caudal vertebrae of Tanystrophaeus (sic!) from thè
Upper Muschelkalk of Bayreuth. However, he ultimately
rejected these latter possibilities.
In 1927 additional remains of Tanystropheus were
found in thè Grenzbitumenzone of Monte San Giorgio
(Cava Tre Fontane, Switzerland). These remains, thought
to belong a single individuai, comprised a few bones
preserved on two plates. Later, in 1929, Bernard Pey-
er’s excavations in thè Grenzbitumenzone of Monte San
Giorgio (Valporina, Switzerland) yielded thè first almost
complete small skeleton of Tanystropheus (neotype, i.e.
specimen PIMUZ T 2791), together with other fragmen-
tary specimens.
In 193 1 Peyer published thè description of all this new
material. On thè basis of thè complete specimen he was
able to identify thè problematic elongate bones of Tanys¬
tropheus conspicuus from thè Upper Muschelkalk of Bay¬
reuth as cervical vertebrae. At thè same time, after a dose
re-examination of thè holotype of Tribelesodon, Peyer
showed that Tribelesodon and Tanystropheus were conge-
neric, and he referred all thè specimens from Besano and
Monte San Giorgio to thè same species: Tanystropheus
longobardicus . Peyer proposed amending thè diagnosis
of thè genus Tanystropheus - up to then based solely on
thè isolated vertebrae of T. conspicuus - and designat-
ing T. longobardicus as thè type species. Moreover, he
suggested designating thè complete skeleton (specimen
PIMUZ T 2791) and thè whole material found in 1929
(sic!) as thè neotype of T. longobardicus. Wild (1974:
148, 151) correctly argued that both proposals had to
be rejected, thè type species of thè genus Tanystropheus
being T. conspicuus by monotypy and thè originai type
being at thè time neither lost or destroyed (irrespective
of thè “neotype” designated by Peyer consisted of more
than one specimen, therefore was per se invalid). In thè
light of thè new discoveries, Peyer gave a re-description
of thè holotype of Tribelesodon. This remains thè best
account of this specimen, which was later destroyed by a
Are caused by thè bombing of thè Museo di Storia Natu¬
rale di Milano during World War II. T. longobardicus was
reconstructed by Peyer as a terrestrial reptile, probably
living dose to thè water. He interpreted thè long neck as
an adaptation towards catching prey from thè shore. Peyer
proposed Tanystropheus as thè unique representative of
thè “Tanysitrachelia” (Peyer, 1931: 89), a new suborder
nested within thè Sauropterygia (this group then includ-
ing Trachelosauria, Nothosauria and Plesiosauria).
In 1974 Rupert Wild reviewed all thè material of T.
longobardicus, including both specimens published by
Peyer in 1 93 1 , as well as undescribed ones resulting from
later excavations conducted by Peyer and Emil Kuhn-
Schnyder. On thè basis of a total of 27 specimens - all but
one housed in thè Collections of thè Palàontologisches
Institut und Museum der Universitàt in Ziirich (PIMUZ) -
Wild (1974) described in detail thè skeletal anatomy of T.
longobardicus, with special emphasis on its skull, which
had remained very poorly known after Peyer’s descrip¬
tion. In addition, he described thè isolated vertebrae of T.
conspicuus from thè German Muschelkalk of Bayreuth.
As thè holotype of Tribelesodon longobardicus (= Tanys¬
tropheus longobardicus ) had been destroyed, Wild could
validly designate specimen PIMUZ T 2791 as thè neo¬
type of T. longobardicus. Wild (1974) gave a detailed
description of thè anatomy, ontogenetic development and
thè adaptation of T. longobardicus. He considered it to
be terrestrial during juvenile stages but predominanti
aquatic as an adult. Wild (1974) regarded Tanystroph¬
eus as a “highly specialized lacertilian”, and included
it in thè Tanysitrachelida (1974: 147), as an infraorder
of thè “Lacertilia”. Later, Wild (1980a) rejected thè
infraorder Tanysitrachelida in favour of thè infraorder
Prolacertiformes, thè latter representing one of thè two
main branches of thè Lacertilia (Wild, 1980a: fìg. 13).
The paper by Wild (1974) remains a benchmark study of
Tanystropheus .
Since then, new species of Tanystropheus have been
described (Jurcsàk, 1975; Wild, 1980a; Rieppel, 2001),
and fragmentary specimens have been referred to thè genus
(Jurcsàk, 1976, 1978, 1982; Vickers-Rich et al., 1999;
Rieppel 2000, 2001; Dalla Vecchia & Avanzini, 2002;
Dalla Vecchia, 2000, 2006; Sulej, 2004; Renesto, 2005),
revealing a western Tethyan distribution and a stratigraphi-
cal occurrence from thè lower Anisian to thè late Norian.
The validity of thè different Tanystropheus species
is summarized below but a full revision of thè genus is
beyond thè scope of this paper.
Wild (1980b: 204; 1987: 39) considered thè possible
synonymy of T. longobardicus and T. conspicuus. In addi¬
tion, he questioned thè validity of T. biharicus (Jurcsàk,
1975), considering this species a probable junior synonym
of T. longobardicus (Wild, 1980a: 12). Fraser & Riep¬
pel (2006) recently erected Amotosaurus rotfeldensis for
specimens from thè Upper Buntsandstein of thè Black
Forest (Germany) previously assigned to “ Tanystropheus ”
antiquus (Ortlam, 1967; Wild, 1980b; Wild & Oostemik,
1984). These authors also provisionally retained thè status
of Tanystropheus antiquus for thè specimens from thè
Lower Muschelkalk of Poland, originally described by
von Huene (1907-1908) but they questioned its validity.
Renesto (2005: 386) questioned thè assignment of T. fossai
(Wild, 1980a) to Tanystropheus, maintaining that thè holo¬
type of T. fossai lacks unequivocal characters justifying its
referrai to that genus. Re-examination of thè holotype of
6
STEFANIA NOSOTTI
T. meridensis (Wild, 1980a) led Fraser and colleagues
(Fraser et al. , 2004; Fraser & Rieppel, 2006: 866) to thè
conclusion that this specimen cannot be distinguished
from thè smallest specimens of T. longobardicus, and they
include it in this taxon. Detailed comparison of thè skull
of T. meridensis with thè new specimens described here
confirms thè statement that T. meridensis and T. longo¬
bardicus should be considered conspecific (Fraser et al.,
in preparation). Renesto (2005) recently described a new
specimen (MCSN 4451) of Tanystropheus from Switzer-
land which, like T. meridensis, was collected in thè Lower
Meride Limestone of Ladinian age. However, due to thè
incomplete preservation of MCSN 4451, together with thè
uncertain status of thè single specimen of T. meridensis
(holotype: Wild, 1980a), Renesto preferred to consider thè
new specimen as T. cf. longobardicus. A preliminary per¬
sonal examination of MCSN 4451, however, did not reveal
striking differences from thè new specimens, suggesting
that it might well be referred to T. longobardicus. Interest-
ingly, Fraser et al. (2004; contra Wild, 1974 and Tschanz,
1988) suggested that there might be two separate taxa
represented in thè material of T. longobardicus from thè
Grenzbitumenzone in thè PIMUZ Collections but further
studies are necessary. At present, T. haasi (Rieppel, 2001)
is thè only unquestioned species of thè genus.
After thè publication of thè monograph by Wild
(1974), several authors discussed, and sometimes drasti-
cally re-interpreted, both thè adaptations of Tanystropheus
(Tschanz, 1985, 1986, 1988; Rieppel, 1989; Taylor, 1989;
Ford, 2002; Renesto, 2005) and its phylogenetic relation-
ships (Benton, 1985; Evans, 1988; Gauthier et al., 1988;
Benton & Alien, 1997; Jalil, 1997; Dilkes, 1998; Peters,
2000a; Rieppel et al., 2003). By contrast, little additional
anatomical investigations have been undertaken, such that
thè descriptions given by Wild (1974; 1980a) remain thè
most complete ever published.
New, remarkably complete and well preserved speci¬
mens of T. longobardicus were collected in thè 1990ies
in sediments outcropping near Besano (Varese Province,
Lombardy, Italy). These specimens, described in this
paper, offer thè first opportunity for new interpretations of
thè anatomy of Tanystropheus since Wild’s (1974) mono¬
graph. The new material also documents thè presence of
Tanystropheus in thè Middle Triassic of Besano. Very
few and incomplete specimens were previously known
from this locality. The holotype of Tribelesodon longo¬
bardicus (see above) is now lost, leaving just two more,
very fragmentary specimens housed in thè PIMUZ Col¬
lections (specimens PIMUZ T 2782 and PIMUZ T 2788,
see Wild, 1974: tab. 1). Three of thè new specimens can
be referred to large-sized individuate, thè first time that
specimens equivalent in size to thè larger specimens in
thè Grenzbitumenzone have been recorded in thè Besano
Formation.
INSTITUTIONAL ABBREVI ATION S
MCSN = Museo Cantonale di Scienze Naturali, Lugano,
Switzerland.
MCSNB = Museo Civico di Scienze Naturali “E. Caffi”,
Bergamo, Italy.
MFSN = Museo Friulano di Storia Naturale, Udine, Italy.
MGB = Museu Geologia de Barcelona, Spain.
MSNM = Museo di Storia Naturale, Milano, Italy.
PIMUZ = Palàontologisches Institut und Museum der
Universitat, Zurich, Switzerland.
YPM = Yale Peabody Museum, New Haven, Connecticut,
USA.
ANATOMICAL ABBREVIATIONS
a = angular
ac = atlas centrum
af = articular facet
afo = adductor fossa
ai = atlas intercentrum
ana = atlas neural arch
arf = articular fossa
art = articular
as = astragalus
at = anterior tubercle
axi = axis intercentrum
bo = basioccipital
bs + ps = basisphenoid-parasphenoid complex
c = cervical vertebra
ca = calcaneum
caf = capitular articular facet
cbl = ceratobranchial I
cc = cristae cranii
cd = caudal vertebra
ce = centrale
eh = Chevron
ci = clavicle
co = coronoid
cor = coracoid
cr = cervical rib
cti = cristae temporales inferiores
d = dorsal vertebra
de = distai carpai
de = dentary
dr = dorsal rib
dt = distai tarsal
e = epipterygoid
eo = exoccipital
f = frontal
fc = fibular condyle
fe = femur
fi = fibula
f-lf = lateral flange of thè frontal
fm = foramen magnum
fp = fossa parietalis
g = gastralia
h = humerus
hs = horizontal shelf
i = ilium
TANYSTROPHEUS LONGOBARDICUS. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
7
icl = interclavicle
iob = impression of thè olfactory bulb
is = ischium
j = jugal
I = lacrimai
lj-pp = lower jaw post-dentary pari
II = lateral lamina of thè surangular
Ilo = lateral lobe of thè main body of thè frontal
lvm = latero-ventral margin
m = maxilla
Mca = Meckelian canal
me = metacarpal
mi = mediai lamina of thè surangular
mio = mediai lobe of thè main body of thè frontal
mt = metatarsal
mth = maxillary tooth/teeth
n = nasal
nc = neural canal
ns = neural spine
of = obturator foramen
op = opisthotic
p = parietal
p-stp = supratemporal process of thè parietal
pa = preatlas
pap = pre-acetabular process
pf = postfrontal
ph = phalanx
pi = palatine
pie = pleurapophysis
pl-mp = maxillary process of thè palatine
plt = palatine tooth/teeth
pm = premaxilla
pmt= premaxillary tooth/teeth
po = postorbitai
pra = prearticular
prf = prefrontal
prf-1 = lobe of thè prefrontal
prf-oa = orbitai aspect of thè prefrontal
prf-pp = palatine process of thè prefrontal
prf-va = postero-ventral aspect of thè prefrontal
prz = prezygapophysis
pt = pterygoid
pz = postzygapophysis
pzc = postzygapophyseal canal
pzp = postzygapophyseal process
pu = pubis
q = quadrate
r = radius
ra = radiale
raf = articular facet for thè rib
s = sacrai vertebra
sa = surangular
sb = sesamoid bone
se = scapula
sep = sclerotic piate
so = supraoccipital
sp = splenial
sq = squamosal
st = supratemporal
stp = stapes
t = tibia
taf = tubercular articular facet
tc = tibial condyle
tp = transverse process
u = ulna
ul = ulnare
v = vomer
vk = ventral keel
vt = vomerine tooth/teeth
MATERIALS
All thè new specimens described in this paper are
housed in thè Paleontological Collections of thè Museo di
Storia Naturale di Milano.
Five specimens come from thè outerops of thè
Besano Formation, or Grenzbitumenzone in thè Swiss
geological literature (Ròhl et al., 2001). On thè basis of
its ammonite fauna, Rieber (1973) referred thè Grenzbi¬
tumenzone to thè uppermost Anisian-lowermost Ladin-
ian. One more specimen was collected in fossiliferous
levels of thè Meride Limestone of Ladinian age, which
is separated from thè Besano Formation by a dolomitic
band (Furrer, 1995).
1) Specimen MSNM BES SC 265, Tanystropheus
longobardicus.
Complete skeleton. Sasso Caldo (SC) quarry, Besano
(BES), Varese Province, Lombardy, northern Italy.
Besano Formation, Lower Ladinian ( curionii Zone),
Middle Triassic.
2) Specimen MSNM BES SC 1018, Tanystropheus
longobardicus.
Incomplete skeleton. Sasso Caldo (SC) quarry, Besano
(BES), Varese Province, Lombardy, northern Italy.
Besano Formation, Lower Ladinian ( curionii Zone),
Middle Triassic.
3) Specimen MSNM V 3663 a b, Tanystropheus
longobardicus, gift by Cesare Ferrario.
Incomplete skull and associated cervical vertebrae
on a pari and counterpart. Spoil from thè Vallone mine,
Besano, Varese Province, Lombardy, northern Italy.
Besano Formation, Anisian-Ladinian boundary. Middle
Triassic.
4) Specimen MSNM BES 215, Tanystropheus cf.
longobardicus.
Isolated dorsal vertebra. Rio Ponticelli quarry, Besano
(BES), Varese Province, Lombardy, northern Italy. Besano
Formation, Anisian-Ladinian boundary, Middle Triassic.
5) Specimen MSNM BES 351, Tanystropheus longo¬
bardicus.
Cervical ribs. Spoil from thè Vallone mine, Besano
(BES), Varese Province, Lombardy, northern Italy. Besano
Formation, Anisian-Ladinian boundary, Middle Triassic.
6) Specimen MSNM V 3730, Tanystropheus cf. longo¬
bardicus, gift by Sergio Rampinelli.
Pes, with partial naturai mould and bone remains of thè
epipodials. Besano, Varese Province, Lombardy, northern
Italy. Meride Limestone, Ladinian, Middle Triassic.
In addition, thè material of Tanystropheus longo¬
bardicus in thè PIMUZ Collections is re-examined.
8
STEFANIA NOSOTTI
METHODS
In thè description of thè materials thè terminology of
Wild (1974) is adopted (unless otherwise stated) in thè
description of processes, foramina and other anatomical
details of each bone. New terms are introduced in quota-
tion marks.
In describing elements of thè pectoral and thè pelvic
girdles, thè terms mediai and lateral are adopted to dif-
ferentiate between thè two main surfaces, except for thè
interclavicle and thè coracoids. The main surfaces of thè
latter elements are referred to as dorsal and ventral.
In describing limb elements, to avoid confusion in thè
terminology relative to their orientation, thè adopted ter¬
minology is specified as follows. The normal position of
thè humerus and thè femur is taken to be directed straight
outwards from thè body, with thè dorsal surface correlated
with thè extensor musculature, and thè ventral surface
correlated with thè flexor musculature. The side facing
anteriorly is referred to as thè anterior side, thè side facing
posteriorly is referred to as thè posterior. As thè precise
orientation of thè epipodial segment remains elusive, thè
normal position of this segment is arbitrarily taken to be
one in which thè epipodials He side by side on a vertical
piane perpendicular to thè median sagittal piane. The sur¬
faces that, in this position, face anterior or posterior are
referred to as thè anterior (extensor musculature) or thè
posterior (flexor musculature) surface of thè bones. The
side closer to thè median sagittal piane is referred to as thè
mediai, thè other side as thè lateral. Finally, thè anatomical
position of thè manus and pes is taken to be one in which
they point forward and are in contact with thè ground.
The surface in contact with thè ground is referred to as thè
palmar and thè piantar respectively (flexor musculature),
thè other one is referred to as thè dorsal (extensor mus¬
culature). As is thè case of thè epipodials, thè manus and
pes have a mediai and a lateral side. In describing move-
ments of thè ankle and thè metatarsals, thè terminology of
Brinkman (1980) is adopted. Flexion of thè ankle is meant
to be a decrease in thè angle at thè ankle, and dorsiflexion
of thè metatarsus is a metatarsal movement associated
with flexion of thè ankle joint; extension of thè ankle is
an increase in thè angle at thè ankle, and plantarflexion of
thè metatarsus is a metatarsal movement associated with
extension of thè ankle.
Skeletal elements in thè different specimens are briefly
described under thè heading of each specimen. An overall
anatomical description, comparisons and discussion con-
ceming thè skull, thè axial and appendicular skeleton are
reported in thè section “Discussion”.
Specimens of T. longobardicus in thè PIMUZ Col-
lections have estimated overall lengths between 53 and
535 cm (Wild, 1974: tab. 1). Wild (1974) considered thè
small specimens to represent juveniles and thè large ones
adults of thè same species, and conjectured that reproduc-
tive maturity corresponded approximately to a size of 2
m total length (on thè basis of presence/absence of post-
cloacal bones and of thè interpretation of thè allometric
growth curves). He considered certain differences in thè
dentition, thè shape of thè bones of thè skull, and in thè
postcranial skeleton to be due to ontogenetic variation.
Recently, Fraser et al. (2004) raised thè possibility that
thè differences observed between thè smallest and thè
larger individuate might instead be related to thè pres-
ence of two separate taxa among thè Grenzbitumenzone
(=Besano Formation) material. Consequently, I will refer
to thè PIMUZ specimens as “small-sized” and “large-
sized” specimens, rather than to juveniles and adults or
even as two distinct species, pending thè formai revision
of taxonomy. Following Wild (1974), I maintain a demar-
cation size of approximately 2 m total length between
small and large individuate, considering that individuate
over 2 m lack distinct tricuspid teeth and dentition on thè
palatine and pterygoid.
See Tab. 1 for thè correspondence of thè letters used
by Wild (1974) to designate thè specimens in thè PIMUZ
Collections with their catalogue number. In thè table, thè
specimens’ overall length is ateo reported.
Table 1 - Correspondence of thè letters used by Wild
(1974) with thè respective catalogue numbers in thè
Tanystropheus longobardicus specimens in thè PIMUZ
Collections, and overall length of each specimen.
(* = estimated). After Wild, 1974: tab. 1.
TANYSTROPHEVS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
9
DESCRIPTION OF THE SPECIMENS
Specimen MSNM BES SC 265
(Figs. 1-9, 49 C-D, 54, 56 A, 58, 65, Pls. I-II, Tabs. 2-5, 8)
The specimen consists of an almost complete, and
mostly articulated skeleton (Fig. 1, Pls. I-II), lying on its
left lateral side. The skull is separated from thè vertebral
column, and is exposed in ventral view. The vertebral
column is exposed in right lateral view, with some disar-
ticulation and rotation in thè sacrai and proximal-caudal
regions. Where disarticulated, thè ribs are stili in dose
proximity to thè corresponding vertebrae. Gastral ribs
cluster in thè trunk region. The elements of thè pectoral
and pelvic girdles are disarticulated but stili lie in an
arrangement indicating their originai position. The fore-
Fig. 1 - Tanystropheus longobardicus, MSNM BES SC 265. Scale bar 50 mm. Preparation: Sergio Rampinelli. Photo: Luciano
Spezia.
10
STEFANIA NOSOTTI
limbs lie side by side ventral to thè trunk, thè right fore-
limb overlapping thè left one. The right hindlimb has been
raised as a consequence of thè twisting of thè vertebral
column, while thè left hindlimb lies ventral to thè trunk.
The estimated overall length of thè specimen is 1 10 cm.
Skull
(Figs. 2-3, 49 C-D)
The complete skull is preserved in ventral view. Its
length, measured from thè tip of thè left lower jaw (=esti-
mated anterior extent of thè premaxilla) to thè posterior
margin of thè parietals, is 5.4 cm. As thè parietals are dis-
placed posteriorly, a more reliable estimate of thè length
of thè skull might be thè distance between thè tip of thè
left lower jaw and thè maximum posterior extent of thè
left squamosal, which is 5.1 cm.
The cranial elements are heavily crushed and elements
of thè neurocranium are randomly displaced posteriorly,
thereby exposing thè ventral surface of thè parietals. The
lower jaws rotated as a unit, with their anterior ends dis¬
placed to thè right, and thè posterior ends to thè left, rela¬
tive to thè sagittal piane. Almost all thè teeth of thè upper
jaws are stili in situ, while thè majority of thè mandibular
teeth are missing.
Premaxilla
The right premaxilla and part of thè left one are concealed
by thè lower jaws. The area of contact between thè right
premaxilla and maxilla is not exposed. The left premaxilla
stili contacts thè left maxilla but thè suture between thè two
elements cannot be distinguished. Consequently, thè number
of premaxillary teeth cannot be ascertained. As in specimen
MSNM BES SC 1018 thè premaxilla bears six teeth (p. 21),
I infer that thè complete premaxillary dentition is seen on thè
left premaxilla of MSNM BES SC 265. The teeth are coni-
cal and have finely striated enamel. The tooth implantation
is subthecodont ( sensu Romer, 1956; Edmund, 1979; Wild,
1974; Motani, 1997; Zaher & Rieppel, 1999).
Maxilla
Both maxillae are preserved. The right maxilla is
almost completely exposed in ventro-medial view, cov-
ered by thè right lower jaw only very anteriorly. Only part
of thè jugal process is missing. The left maxilla is com¬
pletely preserved and exposed in mediai view but it no
longer contacts thè jugal. Almost all thè ankylosed maxil-
lary teeth, and/or thè replacement ones, are preserved in
situ on both maxillae. There are ten teeth in situ in thè
left maxilla and two empty alveoli anterior to them: thè
complete maxillary dentition then includes 12 teeth. The
fìrst unequivocally tricuspid tooth on thè left maxilla is
thè seventh. The sixth has a distinct posterior cusp but
only a very rudimentary anterior one. The fifth only has
at best very rudimentary posterior cusp. Twelve teeth are
also preserved on thè right maxilla. In this case thè fìrst
unequivocally tricuspid tooth is thè fourth.
The raised linguai margins of thè maxillae, dorsal to
thè teeth, should frame thè choanae. The latter, however,
cannot be clearly identified in thè dermal palate.
Vomer
The vomers are entirely concealed by thè lower jaws
but in between thè latter, some isolated vomerine teeth
are exposed. They are very tiny and conical, with striated
enamel.
Palatine
Both palatines are partially exposed. However,
because of thè poor preservation, no details can be dis-
cemed on left palatine. The maxillary process of thè right
palatine is clearly visible but it no longer contacts thè
maxilla. It shows a notched antero-lateral margin fram-
Fig. 2 - Tanystropheus longobardicus , MSNM BES SC 265, skull. Scale bar 10 mm. Photo: Luciano Spezia.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
11
ing thè choana postero-medially, and a weakly concave
postero-lateral margin that framed thè suborbitai fenestra
antero-medially. A conical tooth preserved on thè left side
is tentatively interpreted as a single palatine tooth.
Pterygoid
The right pterygoid exhibits thè transverse process
emerging dose to thè right lower jaw as a stout element,
no longer contacting thè ectopterygoid. Remnants of thè
quadrate process are possibly preserved further posteri-
orly, lateral to thè parietal, and facing thè right quadrate.
No tooth-attachment sites can be identified. The left
pterygoid could not be identified.
Ectopterygoid
No ectopterygoids could be identified in MSNM BES
SC 265.
Frontal
Both frontals are partly exposed in ventral view, roof-
ing thè orbits with their horizontal lateral flanges (pp.
48-49).
plt? prf-pp
Fig. 3 - Tanystropheus longobardicus, MSNM BES SC 265, skull. Drawing and pendi: Fabio Fogliazza.
12
STEFANIA NOSOTTI
Parietal
Both parietals are broadly exposed in ventral view,
and only partly concealed by thè overlapping mandi-
bles. Posteriorly, they are separated along their suturai
contact. The morphological features of thè ventral sur-
face of thè parietal piate that were described by Wild
(1974: 11, fig. 2) can be clearly identifìed. A mediai,
narrow fossa parietalis is framed by thè cristae cranii.
They extend laterally and anteriorly into thè cristae
tempora les inferiores. These in turn form thè thickened
lateral margin of thè parietals. Posterior supratemporal
processes project laterally at an angle of approximately
145°- 150°. They bear an elongate facet for thè reception
of thè squamosals and perhaps thè supratemporals (see
below).
Supratemporal
Paired small bony bars which overlap thè parietals
posteriorly might represent thè supratemporals. However,
because they are displaced, their relationships with thè
neighbouring bones are unclear and their identification
remains equivocai.
Prefrontal
On thè right and left side of thè skull there are two
protuberances which are here interpreted as thè palatine
processes of thè prefrontals.
Squamosal
Both squamosals are preserved. The right squa¬
mosal is partly concealed by overlapping bones. The
left squamosal is fully exposed. Anteriorly, it forms a
sharply pointed postorbitai process which is in dose
proximity to thè jugal. Medially and posteriorly, thè
left squamosal is very broad, probably because of
compression, and a parietal or supratemporal process
cannot be clearly identifìed. The left squamosal does
not meet thè supratemporal process of thè parietal but
this is here considered to be an artefact of preserva-
tion.
The shape of thè squamosal in MSNM BES SC
265 can be described as follows. The bone forms a
“posterior ramus” projecting laterally from thè skull
roof. Then it sharply bends antero-laterally, forming
an “anterior ramus”. Where thè bone bends, a ventrally
projecting triangular lamina forms thè quadrate process.
At approximately two thirds of its length, thè anterior
ramus of thè squamosal bends medially and tapers into
thè postorbitai process. The ventral side of thè squa¬
mosal is grooved.
Postorbitai
No postorbitals could be identifìed in MSNM BES
SC 265.
Jugal
Both jugals are preserved. The left one, exposed in
lateral view, offers more anatomical detail.
The jugal has a triradiate shape, with a long
and slender suborbitai process (=maxillary process
sensi i Wild, 1974: 11) and two posterior processes:
a slender, free-ending quadratojugal process extend-
ing posteriorly, and a broad, long postorbitai process
extending dorsally. The ventral margin of thè jugal is
strongly thickened, forming a ridge along thè lateral
surface of thè bone. The lateral surface of thè pos-
torbital process exhibits an elongate, dorso-ventrally
oriented groove.
Quadrate
The right quadrate is badly crushed and few details
can be discerned. It is considered to be exposed in lat¬
eral view, with thè articular condyle facing thè lower
jaw and thè cephalic condyle facing thè quadrate
process of thè squamosal. Only remnants of thè left
quadrate were tentatively identifìed on thè left side of
thè skull.
Epipterygoid
An isolated, slender bony rod lying posterior to thè
elements of thè neurocranium is interpreted as an epip¬
terygoid (Wild, 1974: 14, figs. 1, 8 b). Its presumed ven¬
tral end is broken but clearly expanded.
Ceratobranchials
Lateral to thè left exoccipital is a ceratobranchial I
(Wild, 1974: figs. 21, 87). The expanded proximal end of
thè bone is directed laterally.
Neurocranium
Elements of thè neurocranium are randomly displaced
posterior to thè parietals. As most of them are badly
crushed and distorted, their interpretation must remain
equivocai.
Both exoccipitals, fused to thè partially preserved
opisthotics, are exposed just posterior to thè parietals, and
stili frame a very deformed foramen magnum.
The very poorly preserved supraoccipital lies further
posteriorly, probably in occipital view.
The basisphenoid-parasphenoid complex is rather
well preserved in ventro-lateral view. The cultriform
process, projecting from thè sphenoidal piate anteriorly,
is distinct. The right alar process and thè anterior part
of thè basisphenoidal piate were pushed outwards. The
basipterygoid process is visible on both sides of thè
basicranium.
Posterior to thè basisphenoid-parasphenoid complex
lies a severely compressed and distorted element, which
was tentatively identifìed as thè basioccipital.
Lower jaw
The mandibular rami separated at thè symphysis
and shifted towards thè median sagittal piane, so that
they are both exposed in lateral view. The right man¬
dibular ramus was probably stretched during fossiliza-
tion, because it is a little longer than thè left one. The
mental foramina open at thè bottom of small pits which
are arranged in a series on thè lateral side of thè den-
taries. The dentaries stili bear some conical and sharply
pointed teeth at a variety of developmental stages. The
presence of poorly distinct cusps on thè posteriormost
preserved tooth on thè left lower jaw remains equivo¬
cai. The dorsally convex suture between thè dentary
and surangular is present on both mandibular rami. The
same is true for thè sutures defining thè articular; on thè
left side thè prearticular is also identifiable, ventral to
thè articular. The presence of a coronoid remains uncer-
tain (p. 61, Fig. 49).
TANYSTROPHEUS LONGOBARDICUS-, RE-INTERPRHTATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
13
Axial skeleton
(Figs. 1, 4-8, 54, 56 A, 58, Pls. I-II, Tabs. 2-3)
Cervical vertebrae and ribs (Figs. 4, 54, 56 A, 58, Pls. I-II,
Tab. 2)
Twelve cervical vertebrae are preserved in right lat-
eral view. Vertebrae two through eleven are complete.
Only remnants of thè atlas are preserved. Part of its right
neural arch, is stili articulated with thè axis (Fig. 56 A).
Two elements posterior to thè parietals are tentatively
interpreted as thè remnants of thè left atlas neural arch
and thè articulated proatlas (Fig. 3). These elements,
however, are very poorly preserved. Moreover, thè pre-
sumed proatlas is larger relative to thè axis than thè same
element in MSNM BES SC 1018 (Fig. 12). The neural
spine on thè 12,h vertebra (Fig. 4) is damaged and it is
overlapped by thè left scapula. Cervical vertebrae three
through ten are deeply grooved because of thè collapse
of their bony walls inside thè hollow vertebral centra
(Wild, 1974: 81).
Cervical vertebrae two through twelve are stili articulated,
except for thè eight and ninth element. At thè joints between
thè 11* and thè 12th cervical vertebrae and that between thè
121*1 cervical vertebra and thè first dorsal, thè cervical verte¬
bral column is bent backwards by almost 180° (Figs. 1, 4,
Pls. I-II). It is no longer articulated with thè skull.
Compression during fossilization makes it impossible
to take reliable measurements of thè vertebrae, apart from
their length (Tab. 2).
The cervical ribs lie dose together forming a thick,
paired bundle that runs parallel to thè cervical column
along thè cervical series seven through twelve (Fig. 1, Pls.
I-II). Anterior to thè seventh cervical, with thè stronger
dorsal bending of thè cervical column, thè rib bundle has
Table 2 - Tanystropheus longobardicus , length of thè cer¬
vical vertebrae in MSNM BES SC 265 and MSNM BES
SC 1018 (* = estimated; n. m. = not measurable).
Fig. 4 - Tanystropheus longobardicus, MSNM BES SC 265, twelfth cervical vertebra and dorsal vertebrae one through three. Water-
color: Massimo Demma.
14
STEFANIA NOSOTTI
become detached from thè vertebrae and projects forward
in a straight line, while thè cervical column is bent back-
wards. However, one of thè ribs of thè fourth pair and all
thè ribs of thè first through third pairs follow thè bending
of thè vertebral column, and He randomly displaced to its
left side. The ribs of thè atlas and thè axis cannot be dis¬
tingui shed from one another.
Dorsal vertebrae and ribs (Figs. 4-5, 7-8, Pls. I-II)
Thirteen dorsal vertebrae are preserved, some of
them very poorly. Dorsal vertebrae one through eleven
are exposed in right lateral view. The first dorsal is more
or less in its originai position relative to thè 12th cervical
(Fig. 4). The latter is tilted backwards, so that thè zyga-
pophyses of thè two vertebrae are slightly shifted relative
to each other. Dorsal vertebrae one through three (Fig. 4)
and fìve through nine (Fig. 7) stili form articulated series.
The fourth dorsal is somewhat displaced relative to thè
preceding and thè subsequent vertebrae. The tenth ver¬
tebra is exceptionally poorly preserved. The 12th and 13th
dorsal vertebrae (“lumbars”, Wild, 1974) are no longer
articulated. The first is exposed in antero-lateral view, thè
second in right lateral view (Fig. 5). The pleurapophysis
( sensu Wild, 1974; see note on p. 69) of thè 12th dorsal is
broken away and lies ventral to thè 13th dorsal. It is not
clear if it is complete. As preserved, it is 12.7 mm long
and 1.3 mm deep.
Fig. 5 - Tanystropheus longobardicus , MSNM BES SC 265, dorsal
vertebrae eleven through thirteen. Watercolor: Massimo Demma.
Measurements of thè centrum length are generally
precluded because of poor preservation. The length of thè
second dorsal vertebral centrum, which is thè best pre¬
served of thè dorsal series, is approximately 1 1 mm and
that of thè 1 1 th is approximately 10 mm.
Few of thè dorsal ribs are preserved. They are disar-
ticulated and scattered but some of them stili He dose to
thè corresponding vertebra. The ribs of thè first and third
pairs are dichocephalous (Fig. 4), while thè remaining
preserved ribs are holocephalous.
Sacrai vertebrae (Figs. 6, 8, Pls. I-1I)
Two sacrai vertebrae are poorly preserved next to thè
right ilium and partly overlapped by thè right femur. They
exhibit stoutly built pleurapophyses ( sensu Wild, 1974;
Fig. 6 - Tanystropheus longobardicus, MSNM BES SC 265, sacrai and
proximal caudal vertebrae. Scale bar 20 mm. Photo: Massimo Demma.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
15
see note on p. 69). They remain articulated together but
completely separate from thè dorsal and thè caudal series.
The centra lie at right angles to thè rest of thè vertebral
column, and are exposed in right lateral view. The pleu-
rapophysis of thè presumed first sacrai is complete, while
that of thè presumed second sacrai is probably truncated.
The distai end of thè complete pleurapophysis is greatly
expanded and bears a wide triangular surface that con-
tacted thè ilium.
Caudal vertebrae (Figs. 1, 6, 8, Pls. I-II, Tab. 3)
The caudal series begins posterior to thè right femur.
It is estimated that there are at least 40 caudals (p. 70).
With thè possible exception of thè vertebral pairs three-
four, four-five, and five-six, thè caudal vertebrae form a
complete articulated series.
Caudals one through three (Figs. 6, 8) are damaged
and deformed. The first caudal is poorly preserved in
right lateral view. The distally truncated right pleu¬
rapophysis (sensu Wild, 1974; see note on p. 69) is
crushed upwards and there is a remnant of thè neural
arch. The centra of thè second and third caudals are
partially rotated around their longitudinal axis and are
exposed in latero-ventral view. The right pleurapophy¬
sis of thè second caudal and both pleurapophyses of
thè third are preserved. The right pleurapophysis of thè
third caudal is 1 1 mm long and 2 mm deep. The right
postzygapophysis of thè second caudal and thè corre-
sponding prezygapophysis of thè third are exposed in
ventral view.
Distai to thè third caudal, thè series is exposed in
right lateral view. The fourth and fìfth caudals (Figs.
6, 8) are largely concealed by thè overlapping bones of
Table 3 - Tanystropheus longobardicus, length of thè
proximal caudal vertebrae in MSNM BES SC 265
(* = estimated; n. m. = not measurable).
thè right hindlimb and thè cervical series. Remnants of
their pleurapophyses project vertically out of thè slab,
indicating that these vertebrae lie on their left side. The
distai end of thè pleurapophyses of these vertebrae was
probably broken away. Indeed there is a fragment of
one such pleurapophyses preserved ventral to thè fìfth
caudal.
Caudals six through eleven (Fig. 6) are very well pre¬
served (thè sixth one is, however, partially overlapped
by thè cervical ribs bundle). Distai to thè llthelement,
thè caudals stili form an articulated series (27.5 cm in
length). The centra are thè only preserved portion of
these vertebrae, and thè articulation between them is
diffìcult to discern.
The measurements of thè proximal caudal vertebrae of
MSNM BES SC 265 are given in Tab. 3.
Gastralia
The gastralia randomly cluster in thè trunk region.
Few of them are preserved.
Appendicular skeleton
(Figs. 1, 7-9, 65, Pls. I-II, Tabs. 4-5, 8)
The most interesting features of thè appendicular skel¬
eton of MSNM BES SC 265 are thè presence of sesamoid
bones in thè elbow and thè knee joints, and thè left tarsus,
whose morphology is discussed on page 72.
Pectoral girdle (Fig. 7)
The elements of thè pectoral girdle are displaced and
only partially preserved and exposed. Only thè right
scapula and thè interclavicle are missing.
The left scapula lies dorsal to thè 1 lth and 12th cervical
vertebrae and is exposed in lateral view. Its short pedun-
cle bears thè articular facet that contributes to thè glenoid
fossa. The antero-dorsal margin of thè bone is heavily
corroded but thè posteriorly projecting process of thè
scapular biade is well preserved.
The coracoids are preserved ventral to thè ante-
riormost dorsal vertebrae, partially overlapping each
other. They are plate-shaped bones, with a short and
stocky glenoid process. A supracoracoideal foramen is
not clearly identifiable. The two coracoids are exposed
in ventral view, because thè articular facet that con¬
tributes to thè glenoid fossa is visible. Thus, thè more
dorsal of thè two is thè left coracoid, shifted across thè
right one.
The clavicles are partially exposed ventral to thè first
dorsal rib. They are rod-shaped, curved elements offering
little anatomical detail. The more ventral element might
be thè mediai part of thè left clavicle, and thè other one
thè lateral part of thè right clavicle (Wild, 1974: 103, figs.
65,91).
Forelimb (Fig. 7, Tabs. 4, 8)
The forelimbs are fairly well preserved. However,
some of their elements are missing and few anatomical
detail of thè preserved bones can be described. They lie
side by side, thè right limb uppermost and overlapping thè
left one. Because of thè displacement of thè bones of thè
pectoral girdle, thè limbs are no longer articulated with
thè latter.
16
STEFANIA NOSOTTI
&
left se
right e!
left cl
left cor
right cor
right h
Fig. 7 — Tanystropheus longobardicus, MSNM BES SC 265, forelimbs. I-IV = Metacarpals. Scale bar 25 mm. Photo: Luciano Spezia.
Pencil: Fabio Fogliazza.
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
17
The right forelimb (Fig. 7) is almost completely
preserved. However, its elements are disarticulated and
displaced relative to one another. The right humerus is
preserved in dorsal view. The epipodials are no longer
articulated with thè humerus and are displaced more or
less as a unit. Given thè relative position of thè radius
and thè ulna, thè radius is preserved in anterior view and
thè ulna, probably, in antero-lateral view. The distai end
of thè ulna is displaced over that of thè radius. The very
poor preservation of thè manus hinders its interpretation
but it seems reasonable to assume that it is preserved in
dorsal view. The manus is approximately in its originai
position relative to thè forearm but no longer articulated
with it. Only two elements of thè carpus, possibly thè
ulnare and thè mediai distai carpai (homology unknown),
are poorly preserved. Differently from metacarpals I-IV,
metacarpal V cannot be reliably identifìed. Metacarpal I
is displaced proximally and no longer aligned with thè
others. By comparison with MSNM BES SC 1018, all
thè phalanges are preserved but they are scattered, and
their proximo-distal originai sequence cannot be reliably
reconstructed.
The left forelimb (Fig. 7) is not complete, because
thè distai ends of thè epipodials and thè manus are not
preserved. The humerus - exposed in dorsal or dorso-
posterior view - lies along thè same longitudinal axis as
thè forearm, and is stili articulated with it. A tiny sesam-
oid bone is identifiable in thè elbow joint. The epipodials
lie side by side exposed in anterior view. The proximal
end of thè ulna overlaps thè radius, and its distai end is
overlapped by thè radius.
The proximal ends of thè humeri have a fiat articular
surface. Distally, thè ulnar condyle is more prominent
than thè radiai one. The epipodials have slightly and
equally expanded proximal ends, with fiat or slightly
convex articular surfaces. The ulnar margin of thè radius
and both margins of thè ulna are concave.
Measurements of thè elements of thè forelimbs in
MSNM BES SC 265 are given in Tab. 4. Length ratios
between different elements of thè forelimbs are given in
Tab. 8.
Table 4 - Tanystropheus longobardicus, measurements of
thè forelimbs in MSNM BES SC 265 (n. m. = not measur-
able).
Pelvic girdle (Fig. 8)
The elements of thè pelvic girdle are displaced and
only partially preserved and exposed.
The elements of thè right side of thè girdle are all pre¬
served and are thicker than thè corresponding elements of
thè left side. They maintain their originai position, having
simply moved away from one another, and are exposed in
lateral view. The ischium, and possibly thè ilium - both
exposed in mediai view - are thè only identifiable ele¬
ments of thè left side of thè girdle.
The right ilium is well preserved, albeit partly con-
cealed by thè large pleurapophysis of thè first sacrai
vertebra. The dorsal iliac biade projects far posteriorly,
ending in a blunt tip. Anteriorly, it is short and rounded
and it is delimited by a supra-acetabular buttress from
thè wide ventral acetabular fossa. As thè acetabular
fossa extends farther anteriorly than thè dorsal biade,
there is a waisted area between thè two parts. A badly
crushed and deformed element ventral to thè 13,h dorsal
vertebra is tentatively interpreted as thè left ilium. Two
shallow fossae might represent facets for thè sacrai pleu-
rapophyses.
The right pubis is largely concealed by thè overlap-
ping left ilium and only its ventralmost portion, antero-
posteriorly expanded, is exposed.
The ischia lie posterior to thè right femur. The right
ischium is partly concealed by thè overlapping caudal
vertebrae. The anterior portion of thè ischiadic piate and
acetabular process is exposed. The left ischium is partly
concealed by thè overlapping right femur. The ventral
portion of thè ischiadic piate and part of thè acetabular
process are exposed.
Hindlimb (Figs. 8-9, Tabs. 5-8)
The hindlimbs of MSNM BES SC 265 are nearly
complete and articulated. The left hindlimb lies ventral
to thè trunk, as expected for a dead animai lying on its
left side. By contrast, as a result of thè twisted sacrai
and proximal-caudal regions of thè vertebral column,
thè right hindlimb has been raised and tumed upside
down. Because of thè displacement of thè bones of thè
pelvic girdle, thè limbs are no longer articulated with
thè latter.
The right hindlimb (Fig. 8) is almost complete but
its elements are partially disarticulated and displaced.
The femur exposes its antero-ventral side. The articu¬
lar surface on its proximal end is grooved. The distai
end displays poorly differentiated condyles, thè tibial
condyle being slightly more prominent than thè fìbu-
lar one. The epipodials are no longer articulated with
either thè femur or thè tarsus. They are preserved in
mediai, slightly anterior view. The proximal end of
thè tibia overlaps that of thè fibula. Distally, thè fibula
was displaced posteriorly. The proximal and thè distai
ends of thè tibia are slightly expanded and convex. The
distai end of thè fìbula is not expanded. The pes is pre¬
served in dorsal view. The only identifiable elements
of thè tarsus are thè calcaneum and thè astragalus (Fig.
9), thè latter poorly preserved and displaced relative to
thè calcaneum. All thè metatarsals are preserved; their
proximal ends are no longer aligned, and metatarsal I is
displaced medially. Metatarsals II through IV and some
phalanges are partially concealed by thè overlapping
cervical vertebral column. By comparison with MSNM
BES SC 1018, all thè phalanges, except thè ungual pha-
lanx of digit live, were identifìed, and thè phalangeal
formula is 2, 3, 4, 5, 4. The phalanges are mostly disar¬
ticulated but their proximo-distal sequences can be stili
identifìed (Fig. 9).
The left hindlimb (Fig. 8) is better preserved than thè
right one. Its elements are all identifiable and only thè
distai phalanges of thè digits are partially displaced. The
femur is exposed in anterior view. The shaft is gently,
sigmoidally curved. The proximal end is expanded and
18
STEFANIA NOSOTTI
Fig. S-Tanystropheus longobardicus, MSNM BES SC 265,hindlimbs. Scale bar 30 mm. Photo: Luciano Spezia. Pencil: Fabio Fogliazza
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
19
weakly convex. The distai end is deflected ventrally.
Only thè tibial condyle is exposed, and its rounded con¬
tour indicates that thè articular surface for thè tibia was
sub-cylindrical in shape. The crus is stili articulated with
thè femur, and rotated on its longitudinal axis together
with thè articulated pes. The crus is thus exposed in
postero-medial view and thè pes in piantar view. A
small sesamoid bone is preserved in thè knee joint. The
tibia has expanded ends, with thè proximal end fiat, and
thè distai end slightly convex. The proximal end of thè
fibula is concealed by thè tibia, but its weakly convex
and not expanded distai end slightly overlaps thè tibia.
The pes is displaced laterally relative to thè crus. The
tarsus (Figs. 9, 65) comprises four ossified elements in
dose juxtaposition: astragalus, calcaneum, distai tarsal
three and distai tarsal four. The astragalus and thè cal¬
caneum meet along a straight line and thè astragalus
slightly overlaps thè calcaneum proximally with a lat-
eral process. A foramen for thè perforating artery (Wild,
1974: 116) cannot be identifìed. The astragalus is a
rather stocky element, elongate transversally relative to
thè longitudinal axis of thè crus. Distally it forms a shal-
low fossa. The calcaneum has a weakly polygonal shape.
Distally, it articulates with metatarsal V. Distally and
medially a line seemingly indicating a contact between
thè calcaneum and distai tarsal four (Fig 65) represents
a crack at thè bottom of a shallow fossa. The proximal
margin of distai tarsal four is not distinct. As preserved,
distai tarsal four partly overlaps thè calcaneum (Fig. 9)
and it is probably a larger element than it appears. Distai
tarsal four contacts metatarsal V laterally and thè proxi¬
mal head of metatarsal IV distally. Distai tarsal three is
displaced and overlaps distai tarsal four. The metatarsals
and thè phalanges are not greatly displaced relative to
one another: metatarsal I through III are displaced proxi¬
mally, and some phalanges are no longer articulated but
their sequence in each digit remains clearly identifiable
(Fig. 9). Metatarsals II through IV, and some phalanges
of thè digits are partially concealed by thè overlapping
cervical vertebral column. The proximal ends of thè
metatarsals overlap, each lateral element overlapping
thè mediai one. By comparison with MSNM BES SC
1018, all thè phalanges are preserved, thè phalangeal
formula being 2, 3, 4, 5, 4.
Measurements of thè elements of thè hindlimbs in
MSNM BES SC 265 are given in Tab. 5. Length ratios
between different elements of thè hindlimbs are given in
Tab. 8.
Fig. 9 - Tanystropheus longobardicus , MSNM BES SC 265, pedes. I-V = metatarsals. Drawing: Fabio Fogliazza.
20
STEFANIA NOSOTTI
Table 5 - Tanystropheus longobardicus , measurements of thè hindlimbs in MSNM BES SC 265 and MSNM
BES SC 1018 (* = estimated; n. m. = not measurable).
Specimen MSNM BES SC 1018
(Figs. 10-29, 49 E, 53 B, 54, 56 B, 59, 62, Pls. III-IV,
Tabs. 2, 5-6, 8)
The specimen consists of a partially articulated and
beautifully preserved skeleton lying on its right lateral
side. The skull is not properly articulated with thè neck,
because thè occipital elements are displaced but lies next
to it. The neck is tilted backwards and is separated from
thè trunk. However, cervical vertebrate two through ten
form an articulated series, except for a separation of thè
fourth from thè fifth. The trunk region comprises a series
of dorsal vertebrae, isolated dorsal ribs, and clustering
gastral ribs. The preserved proximal caudal vertebrae
are scattered on thè slab, while some of thè distai caudals
are articulated and form two isolated series. The pectoral
girdle with both forelimbs, and thè left pelvic girdle with
thè left hindlimb, clearly lie in their originai position rela¬
tive to thè trunk. The right pelvic girdle and hindlimb are
disarticulated, and their elements randomly displaced.
The estimated overall length of thè specimen is 140 cm.
Skull
(Figs. 10-13,49 E, 53 B)
In spite of incomplete preservation, thè skull of
MSNM BES SC 1018 provides some superb anatomi-
cal details, and probably represents thè best example of
thè overall morphology of thè skull in all of small-sized
specimens of T. longobardicus to date. It is exposed in
left ventro- lateral view, so that both mandibular rami
are exposed, thè left one in lateral view, thè right one in
mediai view. The length of thè skull from thè anterior end
of thè left premaxilla to thè posterior end of thè left lower
jaw is 5.8 cm.
The skull is crushed, and thè majority of bones are
displaced to a variable degree. In particular, many of thè
unidentified elements are randomly displaced posteriorly.
By contrast, elements preserved in thè antorbital region
are stili mostly articulated. Dermal bones completing thè
circumorbital series, and forming thè skull roof are only
slightly displaced. Dermal bones of thè palate are partly
exposed in between thè mandibular rami. The lower jaws
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
21
are complete and in situ. Almost all thè teeth of thè upper
and lower lefìt jaws are in situ, while those of thè right side
are mostly displaced.
Premaxilla
Only thè left premaxilla is exposed but it is com¬
plete. It possesses a long and slender maxillary process
contacting thè dorsal margin of thè maxilla, and a short
but clearly delimited nasal process. Posterior to thè
nasal process, thè entire dorsal margin of thè premaxilla
is free, and presumably framed thè extemal naris latero-
ventrally (Figs. 36-37). Tiny foramina open anteriorly
on thè dorsolateral surface of thè premaxilla, possibly
transmitting branches of thè mediai ethmoidal nerve
(Oelrich, 1956).
The premaxilla possesses six teeth - five of which are
preserved in situ - most probably representing thè com¬
plete premaxillary dentition. The teeth are long, conical
and slightly recurved, with fìnely striated enamel. The
third one is thè longest. The premaxillary teeth interlock
with thè anteriormost dentary teeth of thè lower jaw.
Maxilla
Only thè left maxilla is exposed. It is complete, and
stili articulated with thè premaxilla. It is a triangular
bone, bifurcating posteriorly into a long, tapering ven-
tral jugal process, and into a shorter, yet well developed,
dorsal process (processus frontalis, sensu Wild, 1974:
fig. 2). The emarginated region between thè two proc-
esses receives thè lacrimai. Dorsal to thè lacrimai, thè
posterior margin of thè dorsal process contacts thè pre-
frontal. The dorsal margin of thè jugal process no longer
contacts thè jugal. Posterior to thè contact with thè maxil¬
lary process of thè premaxilla, thè dorsal margin of thè
maxilla is free. Originally, it presumably articulated with
thè nasal (p. 46).
Fourteen maxillary teeth are preserved in situ. A gap
between thè two posteriormost teeth suggests that thè
full maxillary tooth count might be of 15. All thè maxil¬
lary teeth are tricuspid. The lateral surface of thè maxilla
exhibits a series of shallow fossae laterally delimited by
bulges. Each of these fossae corresponds to a tooth posi-
tion.
Vomer
Remnants of thè vomer are tentatively identified ante-
rior to thè right palatine but because of very poor pres-
ervation they provide no further detail. However, some
isolated teeth are interpreted as vomerine teeth. They are
tiny and pointed, with striated enamel.
Palatine
Anterior to thè pterygoid, two more dentigerous bones
are interpreted as thè palatines. The element that lies
immediately anterior to thè pterygoid, with four large
alveoli (diameter approximately 1.4 mm), is interpreted
as thè left palatine exposed in ventral view. It stili con¬
tacts thè pterygoid, albeit probably slightly displaced
(thè anteriormost tooth position of thè pterygoid is partly
covered with bone). The maxillary process is apparently
crushed against thè dentigerous region of thè bone. The
element anterior to thè left palatine is interpreted as thè
right palatine exposed in ventral view. It also exhibits four
tooth alveoli approximately as large as those of thè left
palatine. None of thè preserved isolated teeth were identi¬
fied as palatine teeth.
Pterygoid
Only thè left pterygoid is exposed between thè man-
dibular rami, and it is preserved in ventral view. It shows a
well developed transverse process, and a slender, tapering
quadrate process. It bears a row of at least 12 tooth posi-
tions whose diameter becomes smaller posteriorly (from
approximately 0.9 to 0.7 mm). No pterygoid tooth is pre¬
served in situ, and I was not able to single out pterygoid
teeth among thè isolated preserved teeth.
Ectopterygoid and Nasal
No ectopterygoids or nasals could be identified in
MSNM BES SC 1018.
Frontal
Both frontals are well preserved, albeit separated.
The left frontal rotated upwards on its longitudinal
axis, exposing thè ventral side. It exhibits a large axe-
shaped lateral flange, which is medially delimited by
a prominent, sinuously curved ridge. Therefore, thè
flange is concave near thè median sagittal piane, and
deepened into thè cranial cavity. The anterior, lateral,
and posterior margins of thè flange are free. Mediai
to thè lateral flange, thè main body of thè left frontal
extends between thè sinuous margin bordering thè lat¬
eral flange and a straight, yet irregular, mediai margin
originally contacting that of thè right frontal. The ante¬
rior end of thè main body of thè frontal is bifurcate,
with a lateral and mediai lobes interlocking with thè
prefrontal. A notch in thè anterior margin of thè lateral
flange demarcates laterally thè anterior end of thè main
body of thè frontal. Posteriorly, thè left frontal stili
contacts thè parietal but thè latter is badly crushed and
provides no information about thè nature of thè contact
between thè two bones.
The right frontal is displaced into thè orbit as an iso¬
lated element, and it is exposed in dorsal view. Its lateral
flange is partly concealed by thè left frontal and prefron¬
tal. The main body of thè right frontal is thick, and raised
above thè lateral flange. As in thè left frontal, a notch
between thè anterior end of thè main body of thè frontal
and its lateral flange is distinct. A notch in thè postero-
medial end of thè right frontal is interpreted as thè antero-
lateral margin of thè parietal foramen (p. 51).
Parietal
Both parietals are preserved. The left one is crushed
and broken into small pieces although is approximately in
its originai position, posterior to thè left frontal. The right
parietal, exposed in ventral view, is displaced, yet stili in
dose proximity to thè right frontal. Posteriorly it is over-
lapped by thè left postorbital. Its straight lateral margin is
only slightly thickened. Its straight mediai margin ( crista
cranii, sensu Wild, 1974: fig. 2) and oblique antero-medial
margin are thickened. Posteriorly, thè antero-medial
margin is recessed into a subtriangular fossa which bor-
dered thè parietal foramen postero-laterally.
Supratemporal
No supratemporals could be identified in MSNM BES
SC 1018.
22
STEFANIA NOSOTTI
TANYSTROPHEUS LONGOBARDICUS: RH-rNTERPRET ATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
23
Fig. 11 — Tanystropheus longobardicus , MSNM BES SC 1018, skull. Watercolor: Massimo Demma
24
STEFANIA NOSOTTI
Prefrontal
Only thè left, completely preserved, prefrontal is
exposed. The strong compression renders thè interpre-
tation of its complex three-dimensional shape difficult
(p. 52, Fig. 37). I assume that thè posteriormost part of
thè prefrontal rotated upwards on a longitudinal axis
together with thè frontal, thus exposing thè postero-ven-
tral aspect.
The prefrontal is a large bone extending dorso-ven-
trally between thè skull roof and thè lacrimai, and forming
almost entirely thè anterior margin of thè orbit, as well
as its anterior wall. Dorsally, thè prefrontal contacts thè
frontal with a frontal process fìtting into thè biforcate
anterior end of thè main body of thè frontal. Ventrally, thè
prefrontal is slightly shifted relative to thè overlapping
lacrimai, and thè facet receiving thè lacrimai is clearly
visible. Anteriorly, thè prefrontal contacts thè dorsal proc¬
ess of thè maxilla.
Laterally, thè prefrontal develops a sinuous crest dose
to thè orbitai margin, which is dorsally expanded into a
rounded “lobe” (p. 51). In thè fossil, this lobe was raised
and is exposed in ventral view. The dorsalmost part of thè
prefrontal, mediai to thè crest and facing anteriorly, is not
visible.
Nothing can be said about thè presumed contact
of thè prefrontal with thè nasal, because no nasals are
preserved in MSNM BES SC 1018 (but see p. 52 for
discussion).
Lacrimai
Only thè left lacrimai is exposed. It is a small, elongate
subtriangular element. Antero- ventrally it is received into
thè posterior concavity of thè maxilla, and contacts thè
dorsal margin of its jugal process. Dorsally, it is sutured
to thè prefrontal, entering thè anterior margin of thè orbit
to a very limited extent. Posteriorly, thè lacrimai contacts
thè anterior end of thè suborbitai process of thè jugal. A
lacrimai foramen is positioned dose to thè contact of thè
lacrimai with thè maxilla.
Postfrontal
Remains of thè left postfrontal are probably preserved
between thè left parietal and supraorbital process of thè
postorbitai (see below).
Squamosal and Postorbitai
The Identification of thè squamosal and postorbitai in
MSNM BES SC 1018 is problematic.
An arched element forming thè posterior margin of thè
orbit was identified as thè left postorbital. Remnants of
bone projecting posteriorly from thè arch are interpreted
as part of thè postorbital (squamosal process). Conse-
quently, this element appears to be triradiate. The precise
extent of thè antero-dorsal supraorbital process (=orbital
process sensu Wild, 1974: fig. 2) of thè postorbital cannot
be determined, because this area of thè skull is badly
crushed. It is assumed that this area comprises remnants of
thè left parietal, postfrontal and postorbital (supraorbital
process) but thè inter-relationships of thè three elements
cannot be discemed and are only tentatively represented
in Fig. 12. Ventrally, thè postorbital forms a bipartite
jugal process. Each ramus of thè jugal process tapers
into a blunt tip. The anterior ramus of thè jugal process is
in dose proximity to thè groove on thè postorbital proc¬
ess of thè jugal. Postero-dorsally thè postorbital forms a
squamosal process, that is no longer articulated with thè
squamosal. Based on this interpretation of thè postorbital,
thè squamosal would not be preserved in MSNM BES SC
1018.
Alternatively, thè remnants of bone assumed to be
part of thè postorbital (squamosal process) might rep-
resent thè left squamosal, and thè postorbital would be
an arched rather than a triradiate element. I also consid-
ered that an unidentified element preserved dorsal to thè
postorbital might represent a squamosal. However, by
comparison with other specimens of Tanystropheus and
related taxa, these latter interpretations of thè squamosal
and postorbital appear to be less supported (see discus¬
sion on pp. 53-55).
Jugal
Both jugals were identified but thè right one is con-
cealed to a large extent, and only part of its thickened
ventral margin is exposed.
The left jugal is perfectly preserved in lateral view,
closely resembling in shape that described in MSNM
BES SC 265. However, thè jugal of MSNM BES SC
1018 provides more anatomical detail. In particular, thè
longitudinally elongate groove on its postorbital process
is far more distinct and a crest delimits thè groove pos¬
teriorly. Posterior to thè crest a shallow fossa is clearly
visible.
Quadrate
The isolated left quadrate, badly crushed and not com¬
plete, is situated posterior to thè jugal. It is considered to
be in lateral view, with a concave posterior margin and
an expanded dorsal cephalic condyle. The mediai lamina
is partly exposed. The isolated right quadrate was tenta¬
tively identified dose to thè right mandibular ramus but
this element is strongly distorted and does not warrant
description.
Ceratobranchials
Ceratobranchial I lies in dose proximity to thè right
lower jaw. It is very similar in shape to that described for
MSNM BES SC 265.
Neurocranium
Among thè bones of thè neurocranium only thè
supraoccipital was identified. It is exposed as an isolated
element in dorsal view and exhibits a distinct sagittal crest
(-crista occipitali, Wild, 1974: 14).
Scierai plates
Isolated scierai plates are preserved in MSNM
BES SC 1018. Four, with a subrectangular shape, are
positioned dorsal to thè skull. Other elements, mostly
with a very irregular shape and sometimes with a pitted
surface, lie inside thè orbit. They all apparently repre¬
sent crushed and poorly preserved scierai plates. It is
impossible to determine thè full number of thè scierai
plates.
Lower jaw
Both mandibular rami are rather well preserved, thè
left in lateral view and thè right in mediai view. They are
stili dose to each other at thè mandibular symphysis.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
25
Fig. 12 - Tanystropheus longobardicus, MSNM BES SC 1018, skull. Red teeth: isolated maxillary or dentary teeth. The extent of
thè left parietal and postfrontal, and thè dorsal extent of thè left postorbitai (supraorbital process) are arbitrarily chosen. Drawing:
Massimo Demma.
co?
26
STEFANIA NOSOTTI
Fig. 13 - Tanystropheus longobardicus , MSNM BES SC 1018, lower jaws. Drawing: Massimo Demma.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
27
In thè left mandibular ramus (Figs. 13, 49 E) thè
dentary is complete. As previously mentioned, thè
anteriormost conical dentary teeth interlock with thè
premaxillary teeth. The first dentary tooth, preserved in
thè right lower jaw, cannot be seen on thè left side but
four subsequent teeth are distinct. One more conical
tooth is almost completely concealed by thè two pos-
teriormost premaxillary teeth. Thus there was a total
of six conical teeth anteriorly. The tooth immediately
posterior to thè sixth dentary tooth is unequivocally
tricuspid, as are all thè subsequent teeth. These teeth
are overlapped to a large extent by thè corresponding
teeth of thè maxilla but they can be readily counted.
There are eleven tricuspid teeth preserved, and two
empty tooth positions (eighth and llth), giving a total
of 13 tricuspid teeth. The dentary tricuspid teeth are not
interlocking with but linguai to thè maxillary tricuspid
teeth. Shallow fossae delimited by bulges on thè side
of thè dentary mark thè position of each tooth alveolus.
Posterior to thè dentary, thè left mandibular ramus is
crushed and fragmented. The oblique line anteriorly
delimiting thè crushed posterior part of thè lower jaw
probably represents thè suture between thè dentary and
thè surangular (p. 59, Fig. 49). The configuration of
thè postdentary elements is not entirely clear in thè left
lower jaw. Only thè prearticular can be clearly identi-
fied. It has shifted ventrally relative to thè articular.
A facet on thè articular was for thè reception of thè
prearticular. I also tentatively identified part of thè
splenial below thè surangular. It is uncertain whether a
coronoid was present or not (p. 61).
The right mandibular ramus is completely preserved
(Figs. 13, 53 B). The Meckelian canal runs along thè
entire length of thè dentary. The dentary teeth are mostly
displaced. The splenial is a large bone forming to a large
extent thè ventral margin of thè lower jaw. It extends
far anteriorly, roofing part of thè Meckelian canal, and
posteriorly it underlies thè angular and thè articular.
Between thè anterior and posterior slender extensions,
thè splenial forms a large part of thè mediai side of thè
lower jaw. Posteriorly, thè prearticular is clearly identifì-
able as a ribbon-shaped sheet of bone overlapping thè
articular ventrally. The sutures between thè articular,
surangular, angular and thè putative coronoid cannot
be determined, and I can only assume thè approximate
position of these elements. However, thè adductor fossa
bordered by these bones is perfectly preserved, as is thè
articular fossa for thè quadrate.
Axial skeleton
(Figs. 10-12, 14-20, 54, 56 B, 59, Pls. III-IV, Tab. 2)
Cervical vertebrae and ribs (Figs. 10-12, 14, 54, 56 B, 59,
Pls. III-IV, Tab. 2)
Eleven cervical vertebrae are preserved in left lateral
view. Vertebrae two through nine are complete. Some parts
of thè atlas lie as isolated elements dose to thè axis. In
particular, both atlas neural arches are preserved in lateral
view (Figs. 10-12, 56 B). Their slightly dissimilar shape
is probably due to compression. The left atlas neural arch
overlaps thè axis, and it is only slightly displaced relative
to thè axis prezygapophysis. The right neural arch is an
isolated element preserved dose to thè axis. It is clearly
sutured to another element that is tentatively interpreted
as thè proatlas. Two polygonal, articulated elements pre¬
served dorsal to thè left neural arch of thè atlas (Figs. 10-
12) might be thè atlas centrum and intercentrum but this
interpretation remains very doubtful. The tenth vertebra
is incomplete, lacking its posterior half. However, thè
postzygapophysis of thè tenth cervical is stili preserved
in articular contact with thè prezygapophysis of thè llth
(Fig. 14). The llth cervical is completely preserved as an
isolated element. Partial collapsing of thè bony walls into
thè hollow vertebral centra (Wild, 1974: 81) is apparent in
thè cervical series.
Fig. 14 - Tanystropheus Iongobardicus, MSNM BES SC 1018, elev-
enth cervical vertebra in left lateral view. Pz c : postzygapophysis of
thè tenth cervical vertebra. Scale bar 10 mm. Photo: Massimo Demma.
Cervicals two through ten forai an articulated series,
with thè exception of thè fourth and fifth (Fig. 59).
Although thè cervical vertebral column of MSNM BES
SC 1018 is no longer articulated with thè dorsal verte¬
bral column - thè elements of which are mostly isolated
scattered elements - it is clearly tilted backwards, as
commonly observed in other Tanystropheus specimens.
Although thè cervical series is not properly articulated
with thè skull, it maintains its approximate originai posi¬
tion relative to it.
Because of compression it is impossible to take reli-
able measurements of thè vertebrae, apart from their
length (Tab. 2).
The third through tenth pairs of cervical ribs are pre¬
served in dose proximity to thè corresponding centra,
sometimes stili in articulation. Their shafts are exten-
sively broken. Three anteriormost shorter elements dis¬
placed ventral to thè axis were identified as thè ribs of thè
first two cervical vertebrae. Other rib heads and fragments
of their shafts are scattered on thè slab.
Dorsal vertebrae and ribs (Figs. 15-18, 20, Pls. III-IV)
Thirteen dorsal vertebrae are preserved. Fi ve of them,
identified as five through nine or six through ten, form a
partially articulated series stili approximately positioned
in thè trunk region (Pls. III-IV, Figs. 15, 20). They are
exposed in left lateral view. Three additional isolated
28
STEFANIA NOSOTTI
Fig. 15 - Tanystropheus longobardicus, MSNM BES SC 1018, mid-
dorsal vertebra (see discussion on p. 68) in left lateral view. Scale bar 5
mm. Photo: Massimo Demma.
Fig. 17 — Tanystropheus longobardicus, MSNM BES SC 1018, pre-
sumed fourth dorsal vertebra (see discussion on p. 69) in right lateral
view. Scale bar 5 mm. Photo: Massimo Demma.
Fig. 16 - Tanystropheus longobardicus, MSNM BES SC 1018, anteri-
ormost dorsal vertebra (see discussion on pp. 67-68) in left lateral view.
Scale bar 10 mm. Photo: Massimo Demma.
dorsal vertebrae are tentatively interpreted as one of thè
first three anteriormost dorsal, thè fourth, and thè llth.
The anteriormost dorsal (Fig. 16) is exposed in left lateral
view: its centrum is 14.8 mm long. Both pre- and postzyg-
apophyses and thè posterior end of thè neural spine are
broken. The putative fourth dorsal (Fig. 17) is exposed in
right lateral view: its centrum is 15.4 mm long. A fracture
crosses thè posterior area of thè neural arch and continues
along thè border of thè posterior end of thè centrum. The
putative llth dorsal (Fig. 18) is exposed in right lateral
view: its centrum is 14.6 mm long.
Fig. 18 - Tanystropheus longobardicus , MSNM BES SC 1018, pre-
sumed eleventh dorsal vertebra (see discussion on p. 69) in right lateral
view. Scale bar 5 mm. Photo: Massimo Demma.
The preserved, disarticulated dorsal ribs mostly cluster
in thè trunk region. They are stout, longitudinally grooved
elements. All of them are holocephalous. A single, short
dichocephalous rib is preserved close to thè isolated right
femur (Pls. III-IV). It was clearly associated with one of
thè anteriormost dorsals.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
29
Fig. 19 - Tanystropheus longobardicus, MSNM BES SC 1018,
sacrai vertebra in right lateral view. Scale bar 5 mm. Photo: Massimo
Demma.
Sacrai vertebrae (Fig. 19)
One sacrai vertebra is preserved as an isolateci ele-
ment exposed in right lateral view. It bears a stout, albeit
crushed, pleurapophysis ( sensu Wild, 1974; see note on
p. 69). The centrum is slender and thè low neural spine
raises posteriorly into a rounded process.The amphicoe-
lous centrum of this vertebra is 14 mm long.
Caudal vertebrae (Pls. III-IV)
Twelve caudal vertebrae are scattered on thè slab.
A group of six caudal vertebrae exposed in lateral or
ventro-lateral view is preserved in thè same area as thè
scattered elements of thè right hindlimb. When measur-
able, thè centrum length of these vertebrae is approxi-
mately 12 mm, although thè caudal to thè left of thè
anteriormost preserved dorsal (PI. IV, d,_3?) has a centrum
length of 16.4 mm. The centrum is amphicoelous. The
shape of these vertebrae is difficult to interpret but thè
presence of pleurapophyses (sensu Wild, 1974; see note
on p. 69), sometimes very wide at thè base, indicates that
they are most likely proximal caudals.
An additional proximal caudal vertebra is preserved
in antero-lateral view dose to thè distai part of thè right
tibia. The right pleurapophysis is distally truncated and a
line at thè very base of it might be interpreted as a suture
(see note on p. 69). The orientation of thè prezygapo-
physeal articular facets is sub-horizontal. Ventral to thè
amphicoelous centrum, thè Chevron is preserved. It is a
Y-shaped element and thè two rami are proximally con-
nected by a transverse bony bar.
Three more caudal vertebrae lie in thè area between thè
cervical vertebral column and thè left femur. Two of them
lie to thè right of thè fìrst articulated series of distai caudal
vertebrae. They are very deformed and their shape is difift-
cult to interpret. One of thè two appears to preserve pleura¬
pophyses and on this basis it might be assumed to be part of
thè proximal caudal vertebral column. A third caudal lies to
thè right of cervical ten, and is preserved in left lateral view.
The centrum is 16.9 mm long. By direct comparison with
MSNM BES SC 265, this vertebra is tentatively identified
as thè ninth caudal. A pleurapophysis is lacking but a shal-
low, yet distinct tiny fossa is present on its lateral surface.
This small fossa is seen in other isolated caudals of MSNM
BES SC 1018 ventral to thè pleurapophyses.
Finally, two vertebrae preserved in anterior view are
identified as distai caudal vertebrae. They are very small
and lack pleurapophyses. One is dose to thè mid-dorsal
series in thè trunk region. The second is partially over-
lapped by thè proximal part of thè right tibia and fibula.
Its centrum is weakly amphicoelous; only thè pedicel of
thè neural arch is preserved.
Like MSNM BES SC 265, thè distai caudals form two
articulated series in which thè centra are thè only pre¬
served vertebral elements. The articulation between thè
centra, however, is hardly identifiable. The fìrst series is
5 cm long and is very poorly preserved. The second is 20
cm long and better preserved. Remnants of bone proximal
to thè fìrst series probably represent other distai caudal
vertebrae but they are exceptionally poorly preserved.
Gastralia
The gastralia are preserved in their originai position,
ventral to thè trunk. They are closely packed, and, in spite
of disarticulation of thè different elements, they clearly
form a “skeletal unif ’. The anterior elements are preserved
in ventral view, while thè posterior ones appear to be in
anterior view. It is estimated that thè gastral skeleton
includes about 30 units, each composed by four elements.
Appendicular skeleton
(Figs. 20-29, 62, Pls. III-IV, Tabs. 5-6, 8-9)
Specimen MSNM BES SC 1018 is unique amongst
all known specimens of Tanystropheus with respect to thè
beautiful preservation of thè limbs. By contrast, thè pre¬
served elements of thè pectoral and thè pelvic girdles are
displaced, and some of them are fragmentary or broken.
Pectoral girdle (Fig. 20)
All thè elements of thè pectoral girdle, with thè pos-
sible exception of thè interclavicle, can be identified.
The left scapula is complete, and exposed in lateral view.
The thick ventral peduncle bears thè articular facet contribut-
ing to thè glenoid cavity. Dorsal to thè peduncle, thè scapular
biade is strongly expanded and fan-shaped, projecting more
posteriorly than anteriorly. It exhibits concentric striation.
The right scapula is badly broken but its overall shape is stili
recognizable. It is preserved in mediai view.
The two coracoids lie side by side, concealed partly by
thè left humerus. They are plate-shaped elements. Their
glenoid area is not exposed. The element to thè left is
probably thè left coracoid in ventral view, and thè one to
thè right is thè right coracoid in dorsal view.
The two clavicles are preserved in between thè scapu-
lae. The element overlapping thè left scapula is most
likely thè left clavicle in ventral view, while thè other
is thè right clavicle in anterior or posterior view (Wild,
1974: 103, figs. 65, 91).
An element overlapped by thè right clavicle is tenta¬
tively interpreted as thè interclavicle. Two distinct proc-
esses which embrace thè clavicles project laterally from
thè anterior piate. This presumed interclavicle has a shape
different from thè only other interclavicle reported in thè
material of Tanystropheus (Wild, 1974: 103, fig. 64).
30
STEFANIA NOSOTTI
Fig. 20 - Tanystropheus longobardi cus, MSNM BES SC 1018, pectoral girdle and mid-dorsal vertebra in left lateral view. Scale bar
15 mm. Photo: Massimo Demma.
Forelimb (Figs. 21-23, Tabs. 6, 8-9)
Both forelimbs are superbly preserved in complete
articulation. They are without doubt thè best preserved
forelimbs of all known Tanystropheus longobardicus
specimens.
The right humerus (Fig. 21) is exposed in anterior
view. Because of thè twisting of thè two articular ends
relative to each other (Wild, 1974: 106, fig. 67), thè
expansion of thè proximal end and thè profile of thè distai
end are evident. Although thè distai end is crushed, thè
rounded shape of thè two condyles is clear. The forearm is
preserved in anterior view, and thè manus in dorsal view.
The radiale and thè lateral distai carpai are each broken
into two pieces.
The left forelimb (Fig. 22) is stretched, thè humerus,
exposed in ventral view, lying along thè same longitudi-
nal axis as thè forearm, exposed in anterior view. The
manus is exposed in dorsal view. Because ot compres-
sion, thè proximal end of thè humerus lies on thè same
piane as thè distai one. A crack crosses thè proximal end
of thè humerus but all other elements are intact. A sesam-
oid bone is preserved in thè elbow joint.
The humeri have a more or less straight shaft and
expanded ends. The proximal end is slightly convex, thè
distai one is differentiated into two condyles.
The epipodials are of sub-equal length, and have
approximately equally expanded ends. In anterior view,
thè proximal end of thè radius slightly overlaps that of thè
ulna, and vice versa distally. The radius is stouter than thè
ulna. Its ulnar margin is concave. Both ends are slightly
expanded and have fiat articular surfaces. The ulna is a
slender element with concave margins and expandend
ends, thè proximal slightly more so than thè distai one.
The articular surfaces are distinctly convex, particularly
thè distai one. A spatium interosseum is present.
The carpus includes four ossifìed elements (Fig. 23).
The proximal two, thè radiale and thè ulnare, meet each
other along a straight line and enclose a well-defìned per-
forating foramen. Following Wild (1974), thè two distai
carpai elements are referred to as thè mediai and thè lat¬
eral one, because their homology is at present unknown.
Contra Wild (1974: 109, fig. 70), thè largest distai carpai
is thè lateral one. It is a roundish element positioned
between thè ulnare, proximally, and thè proximal heads
of metacarpals III and IV, distally. Medially, thè lateral
distai carpai contacts a rounded mediai distai carpai. The
latter is in juxtaposition with thè proximal heads of meta¬
carpals Il and III. There is a wide gap between thè mediai
distai carpai and thè radiale, and thè mediai region of thè
carpus is unossified.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
31
■ ■ S
'•*? J is '• •*
Fig. 21 - Tanystropheus longobardicus, MSNM BES SC 1018, right forelimb. Scale bar 20 mm. Photo: Luciano Spezia
32
STEFANIA NOSOTTl
Fig. 22 - Tanystropheus longobardicus, MSNM BES SC 1018, left forelimb. Scale bar 20 mm. Photo: Luciano Spezia.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRET ATION S OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
33
Fig. 23 - Tanystropheus longobardicus, MSNM BES SC 1018, left
manus in dorsal view. Broken lines indicate thè contours of thè caudal
vertebra which was erased to highligth thè carpus. I-V = metacarpals.
Scale bar 5 mm. Photo: Massimo Demma.
Metacarpals I and V are far shorter than thè others,
with V being thè shortest. Metacarpals II through IV are
distinctly longer. Metacarpal III is thè longest, followed
by IV and then II. The proximal ends of metacarpals
II through IV overlap. In dorsal view each mediai ele-
ment overlaps thè lateral one. The phalanges do not
Ìlook as thick and shortened as in Wild’s reconstruction
(1974: fig. 70). The phalangeal formula of thè manus is
2, 3, 4, 4, 3 (see Tab. 9 for comparison with other protoro-
saurian taxa).
Measurements of thè elements of thè forelimbs in
MSNM BES SC 1018 are given in Tab. 6. Length ratios
between different elements of thè forelimbs are given in
Tab. 8.
Table 6 - Tanystropheus longobardicus, measurements
of thè forelimbs in MSNM BES SC 1018 (* = estimated;
n. m. = not measurable).
Pelvic girdle (Figs. 24-25, Pls. III-IV)
The left elements of thè pelvic girdle (Fig. 24) are
better preserved than thè right ones, albeit fragmentary.
The ilium, ischium and pubis approximately maintain
their position relative to one another but have separated.
They are exposed in lateral view. Fragments of thè right
elements of thè pelvis are scattered in thè same area.
The left ilium has a posteriorly projecting dorsal biade
and a wide, ventral acetabular portion. The ilium contrib-
utes most to thè acetabulum. A distinct waist separates
thè two parts. The dorsal biade tapers posteriorly into a
blunt process. Anteriorly, it is concealed to a large extent
by a fragment of thè supposed right pubis but it is clearly
far shorter than posteriorly. A short pre-acetabular proc-
34
STEFANIA NOSOTTI
Fig. 24 - Tanystropheus longobardicus, MSNM BES SC 1018, pelvic girdle. Scale bar 15 mm. Photo: Massimo Demma.
ess or tubercle is present anteriorly. The posterior part of
thè dorsal biade deepens into a fossa, dorsally sulcated by
a groove. The acetabular fossa is dorsally framed by thè
acetabular buttress. Its ventral margin is not identifiable
but posterior to it thè articular facet for thè ischium is dis-
tinct. The right ilium (Pls. III-IV) is displaced dose to thè
1 l,h cervical vertebra, and is exposed in lateral view. Only
its post-acetabular portion is preserved. The iliac biade
has thè same shape as thè left side.
The left pubis is complete. It is differentiated into an
acetabular part and a ventral biade. The latter is antero-
posteriorly expanded and its surtace exhibits concentric
striation. In thè acetabular region, a distinct articular
peduncle for thè ischium projects posteriorly. Conse-
quently, thè posterior margin of thè pubis, bordering
thè thyroid fenestra, is strongly concave. A wide, ovai
obturator foramen pierces thè pubis in this area. Ante-
rior to thè peduncle, thè acetabular facet of thè pubis
is distinct. Anterior to thè acetabular facet, thè margin
of thè pubis is damaged. In this area it contacted thè
pre-acetabular part of thè ilium and apparently formed
a pubic tuberosity. Contra Wild (1974: 112, fig. 71), a
processus lateralis similar to that observed in thè extant
squamates was not identified in thè pubis ot MSNM
BES SC 1018. Dorsally, thè anterior margin of thè pubis
is straight, ventrally, is very slightly convex. A fragment
of bone overlapping thè anterior part of thè iliac biade is
tentatively interpreted as thè proximal part of thè right
pubis.
Only thè fan-shaped, ventralmost part of thè left
ischium is preserved, broken in two pieces. Fragments of
thè same region of thè right ischium are preserved dose to
thè left pubis. The surface of thè ischiadic blades displays
concentric striation.
The left side of thè pelvic girdle of MSNM BES SC
1018 is reconstructed in Fig. 25.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
35
Fig. 25 - Tanystropheus longobardicus , reconstruction of thè pelvic
girdle (left side in lateral view) based on MSNM BES SC 1018. Water-
color: Massimo Demma.
Hindlimb (Figs. 26-29, Pls. III-IV, Tabs. 5, 8-9)
The preserved elements of thè right hindlimb (Fig.
26, Pls. III-IV) are disarticulated and scattered. The right
femur is preserved in antero-dorsal view. The shaft is prox-
imally broken, at approximately four fifths of thè length
of thè bone. Another piece lies to thè right of this larger
piece. The isolated proximal end of thè femur (Fig. 26)
lies ventral to thè trunk. The articular surface is convex.
The distai end is deflected ventrally, giving thè shaft a
gently curved shape. The tibial condyle widely conceals
that for thè fibula. The rounded shape of thè condyles sug-
Fig. 26 - Tanystropheus longobardicus , MSNM BES SC 1018, proxi¬
mal end of thè right femur. Scale bar 5 mm. Photo: Massimo Demma.
gests that they formed sub-cylindrical articular surfaces
for thè tibia. The right tibia is broken into two pieces. One
represents approximately two thirds of thè overall length
of thè bone and includes thè proximal end. It is displaced
to a position dorsal to thè trunk, and is exposed probably
in dorsal view. The crushed distai end with thè distalmost
part of thè shaft is preserved on thè other side of thè trunk.
Both ends of thè tibia are expanded. The proximal end of
thè right fibula with thè proximalmost part of its shaft is
stili articulated with thè tibia. The larger part of thè shaft
with its distai end lies dose to thè femoral head. Disartic¬
ulated elements of thè right pes are preserved in thè same
area as thè femoral shaft. Distai tarsal three and distai
tarsal four together with metatarsals I through IV lie dose
to one another, and there are some scattered phalanges.
The astragalus and calcaneum are thè only articulated ele¬
ments of thè right pes. The astragalus and calcaneum unit
is exposed in plantar-distal view. The mediai swollen part
of thè astragalus was crushed on its lateral part, exposing
its distai surface. Consequently, thè astragalus is strongly
deformed.
All thè elements of thè left hindlimb (Fig. 27) are pre¬
served to thè right side of thè neck. Although not properly
articulated, thè femur, crus, and pes are preserved dose
to one another in their originai proximo-distal sequence.
The left femur is exposed in antero-dorsal view. Its proxi¬
mal head is broken into small pieces but it is stili clearly
facing thè acetabular part of thè left ilium. The shaft is
gently, sigmoidally curved. The two rounded distai con¬
dyles are exposed, thè tibial widely concealing that for thè
fibula (Fig. 29). The tibia and thè fibula rotated slightly
clockwise on their longitudinal axis and fell as a unit. The
two bones lie thus parallel to each other and are probably
exposed in antero-medial view. In this view, thè proximal
end of thè tibia is more expanded than thè distai one. The
mediai margin of its shaft is slightly concave. The fìbula
is a slender bone with a sigmoidal shape and no expanded
ends. The tibia and thè fibula are equal in length. The pes
(Fig. 28) is exposed in piantar view. All thè elements of
thè tarsus are preserved (Fig. 62) but thè articulated astra¬
galus and calcaneum unit was displaced and tumed upside
down, so exposing its dorsal side. The astragalus and
calcaneum meet along a straight line and enclose a well-
defined foramen for thè perforating artery (Wild, 1974:
116). The articular facet for thè fibula formed by thè two
bones is exposed on thè proximal side of thè astragalus
and calcaneum unit. On thè distai side, both astragalus and
calcaneum contribute to a fossa, proximally delimited by
a sharp margin. The mediai part of thè astragalus is partly
overlapped by metatarsal V. Distai tarsal three and distai
tarsal four contact metatarsals III and IV respectively but
distai tarsal four does not meet thè proximal end of meta¬
tarsal V. Therefore some dislocation probably occurred.
The metatarsals and thè phalanges are completely pre¬
served in articulation. The proximal ends of thè metatar¬
sals overlap, each lateral element overlapping thè mediai
one. A small but clearly delimited ventral tubercle is seen
dose to thè lateral margin of metatarsal V. The phalangeal
formula of thè pes is 2, 3,4, 5,4 (see Tab. 9 for a comparison
with other protorosaurian taxa).
Measurements of thè elements of thè left hindlimb in
MSNM BES SC 1018 are given in Tab. 5. Length ratios
between different elements of thè hindlimbs are given in
Tab. 8.
36
STEFANIA NOSOTTI
Fig. 27 - Tanvstropheus longobardicus , MSNM BES SC 1018, left hindlimb. Scale bar 30 mm. Photo: Luciano Spezia
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
37
Fig. 28 - Tanystropheus longobardieus, MSNM BES SC 1018, left pes
I-V — metatarsals. Scale bar 10 mm. Photo: Roberto Appiani.
in piantar view. Astragalus and calcaneum in dorsal view.
I
38
STEFANIA NOSOTTI
Fig. 29 - Tanystropheus ìongobardicus , MSNM BES SC 1018, left
hindlimb, knee joint. Scale bar 10 mm. Photo: Massimo Demma.
Specimen MSNM V 3663 a b
(Figs. 30-32)
Specimen MSNM V 3663 is a skull with five associ¬
ateci cervical vertebrae preserved on a part (V 3663 a), and
a naturai mould with remains of bone and teeth on a coun-
terpart piate (V 3663 b). It is notable for its large size and
thè preservation of some elements of thè temporal region
and of thè vomerine dentition.
By comparison with thè skulls of MSNM BES SC 265
and MSNM BES SC 1018, thè overall length of thè indi¬
viduai represented by this specimen is estimated to have
been at least 3 m. In addition, thè length of thè most com¬
plete cervical vertebra in MSNM V 3663, representing
an element of thè series six through nine, is 17 cm. This
length corresponds to that of thè cervical vertebrae six-
seven in specimens estimated to be approximately 3.5 m
long (PIMUZ T 2790 and PIMUZ T 2819) in thè PIMUZ
Collections (Wild, 1974: tab. 3).
The length of thè skull, measured from thè posterior-
most extent of thè parietals (as preserv ed) to thè anterior-
most extent of thè right lower jaw, is 12 cm. The skull is
exposed in ventral view and very poorly preserved. It is
heavily crushed and thè surface of thè bone is wom away.
Few elements of thè skull could be identified. In thè pre¬
orbitai region there is a small part of thè premaxilla with
conical premaxillary teeth, part of thè right maxilla with
conical maxillary teeth, and thè vomers. The latter exhibit
a series of tooth alveoli: seven on thè right vomer and five
on thè left. Teeth are present in some of these. These teeth
are very small and apparently conical but their tips are
broken. The anteriormost of thè preserved teeth on thè left
vomer has a basai diameter of 1 .3 mm.
Some elements of thè skull roof, circumorbital series
and of thè temporal region are also exposed. Remains
of thè parietals and frontals can be discemed but they
are indistinct. By contrast, thè right postfrontal, stili
articulated to thè fronto-parietal piate, can be clearly
identified. It is articulated with another element which is
interpreted as a fragmentary postorbital. Finally, thè right
jugal is exposed in mediai view. Posterior to thè elements
described, a partial naturai mould and remains of bone
probably represent thè squamosal, and indicate thè poste-
riormost extent of thè skull.
The anteriormost part of thè right dentary, exposed in
lateral view, overlaps thè premaxilla. Another fragment of
thè right lower jaw is preserved just anterior to thè right
jugal. The fragmented, anteriormost part of thè left den¬
tary and a partial naturai mould of thè left lower jaw are
seen on thè left side of thè skull.
On thè counterpart piate, a partial naturai mould of
thè right lower jaw and remains of it can be identified.
Moulds and remains of thè premaxillary and maxillary
teeth are distinct as well.
The preserved cervical vertebrae are crushed and
fragmented and their morphology cannot be described
in detail. It can be only stated that they have thè typical
elongate shape of cervicals three through ten. One pair of
fragmentary, originally articulated vertebrae is preserved
dose to thè skull. Another segment of thè cervical verte-
bral column is represented by three, originally articulated
elements: thè posterior end of one vertebra, a second more
complete vertebra and thè anteriormost part of a third.
Fragments of cervical ribs run ventral to thè vertebrae.
Specimen MSNM BES 215
(Fig. 33)
Specimen MSNM BES 215 is an isolated mid-dorsal
vertebra in posterior view (p. 68). It is significant for its
large size and thè peculiar morphology. Although embed-
ded in a slab, thè vertebra is fairly three-dimensionally
preserved.
The vertebra lacks thè dorsal part of thè neural spine.
Its height, measured from thè ventral surface of thè cen-
trum to thè maximum dorsal extent of thè broken neural
spine, is 7.4 cm. The height of thè centrum is approxi¬
mately 3 cm. By comparison with thè height of thè cen¬
trum of thè mid-dorsal vertebrae of MSNM BES SC 1018
and thè large T. conspicuus (Wild, 1974: fig. 54), thè
overall size of thè individuai to which MSNM BES 215
belonged is estimated to have been at least 5 m.
The centrum has an elliptic dorso-ventrally elongate
shape. Its articular surface is wom away but a thickened
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPEC1MENS FROM BESANO
39
Fig. 30 - Tartystropheus longobardicus, MSNM V 3663 a, piate. Scale bar 1 00 mm. Preparation: Fabio Fogliazza. Photo: Luciano Spezia.
margin and a shallow centrai fossa are identifiable. Short
and stocky transverse processes project laterally from
thè area between thè centrimi and thè very base of thè
neural arch. The neural canal is pear-shaped and filled
with matrix. Above thè canal there are short postzyga-
pophyses with fiat articular surfaces that slightly slant in
a dorso-lateral/medio-ventral piane. These wide articular
surfaces reach thè area just above thè neural canal. A short
bony shelf develops above thè neural canal, and forms thè
floor of a very shallow postzygapophyseal trough ( sensu
Rieppel, 2001), within which paired postzygapophyseal
canals {sensu Rieppel, 2001: fìgs. 2-3) can be seen,
albeit filled with matrix. This latter feature is interesting
because postzygapophyseal canals were never previously
described in thè post-cervical vertebrae of Tanystropheus
(p. 68). For this reason MSNM BES 215 is here consid-
ered as Tanystropheus cf. longobardicus, albeit on thè
basis of its stratigraphical position it could be ascribed to
T. longobardicus.
The neural spine of MSNM BES 215 is broken dor-
sally. A median longitudinal ridge runs along it, and
diverges ventrally above thè neural canal and between thè
fiat articular surfaces of thè postzygapophyses, framing
thè postzygapophyseal canals medially. The dorsal liga-
ment would have been inserted on this ridge (Wild, 1974:
fig. 38 e).
40
STEFANIA NOSOTTI
Fig. 31 - Tanystropheus longobardicus, MSNM V 3663 a, piate, skull. Scale bar 20 mm. Photo: Roberto Appiani. Drawing: Massimo
Demma.
i
TA NYSTROPHE US LONGOBA RDIC US : RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
41
Fig. 32 - Tanystropheus longobardicus, MSNM V 3663 b, counterplate. Scale bar 10 mm. Photo: Luciano Spezia.
Fig. 33 - Tanystropheus cf. longobardicus, MSNM BES 215, mid-dorsal
vertebra in posterior view. Scale bar 10 mm. Photo: Luciano Spezia.
Specimen MSNM BES 351
(Fig. 34)
Specimen MSNM BES 351 comprises isolateci frag-
ments of thè shafts of cervical ribs belonging to an
individuai larger in size than MSNM BES SC 265 and
MSNM BES SC 1018. This can be inferred from thè
diameter of thè shafts which is approximately 2 mm in
MSNM BES 351, and 0.5-1.0 mm in MSNM BES SC
265 and MSNM BES SC 1018. The diameters of thè cer¬
vical ribs in specimen PIMUZ T 2819 (estimated overall
Table 7 - Tanystropheus cf. longobardicus , measurements
of MSNM V 3730.
42
STEFANIA NOSOTTI
Fig. 34 - Tanystropheus longobardicus , MSNM BES 351, fragments of cervical ribs. Scale bar 30 mm. Photo: Luciano Spezia.
length 3.65 m, Wild, 1974: tab. l)are 1 .0-2.0 mm proxi-
mally and thickening to a maximum between 2.0 and 3.0
mm (Wild, 1974: 60, tab. 4).
Specimen MSNM V 3730
(Figs. 35, 61, Tab. 7)
Specimen MSNM V 3730 is an isolated pes with all
thè elements preserved, and mostly in articulation. A
partial naturai mould and remains of thè disarticulated
epipodials are also present but are poorly preserved and
provide no information on their detailed morphology.
On thè basis of thè phalanges, which bear longitudinal
grooves along their shafts and prominent distai condyles
separated by a groove, this specimen most probably rep-
resents a right pes in piantar view. By comparison with
MSNM BES SC 1018, thè overall size of thè individuai
to which thè pes belonged is estimated to have been
approximately 1.50 m.
The tarsus of MSNM V 3730 is unique for all known
Tanystropheus material in preserving all thè elements in
their originai position and articulation. The calcaneum
is a polygonal element with roundish and thickened
lateral margin. Its concave distai margin matches thè
rounded proximal head of metatarsal V. The astragalus is
a proximo-distally elongate element, obliquely oriented
within thè tarsus. Its proximal end meets thè calcaneum
in a straight line. Half way along it there is an elon¬
gate foramen for thè passage of thè perforating artery
(Wild, 1974: 116). Distally, thè astragalus expands into
a rounded head, that does not contact metatarsal II. The
distai ossifìcations of thè tarsus are roundish elements.
and distai tarsal three is far smaller than distai tarsal four.
Distai tarsal three lies mediai to thè distai head of thè
astragalus and is in dose juxtaposition with thè proximal
end of metatarsal III. Distai tarsal four lies in thè lateral
embayment of thè astragalus and contacts thè proximal
head of metatarsal IV distally. It contacts distai tarsal
three medially and articulates with thè proximal rounded
head of metatarsal V laterally.
The metatarsals are tightly packed, and their proximal
ends overlap. On thè proximal end of metatarsal IV there is
a distinct facet for thè reception of metatarsal V. The bony
wall of metatarsals I through IV is partly collapsed. Lateral
to thè contact with thè calcaneum thè surface of metatar¬
sal V is recessed into a shallow fossa. Distally, metatarsal
V forms a rounded articular head for thè fìrst phalanx of
digit five. Lateral to that there is a distinct rounded proc-
ess, possibly representing a lateral piantar tubercle (p. 73).
An “outer process” with a distinct tuberosity is developed
proximo-laterally (p. 73). The lateral margin of metatarsal
V is concave and thickened.
The metatarsals and thè phalanges of digits one and
two form articulated series. The ungual phalanx of digit
three is displaced, as are thè three articulated distai
phalanges of digit four and thè two articulated distai
phalanges of digit five. Finally, thè fìrst phalanx of digit
five is displaced across thè distai ends of metatarsals III
and IV, and lies at right angles to thè second phalanx of
digit five. Like T. meridensis, MSNM V 3730 was col-
lected in thè Meride Limestone of Landinian age. T meri¬
densis is probably conspecific with T. longobardicus (see
“Introduction”). However, pending thè format revision of
taxonomy, MSNM V 3730 is provisionally considered as
Tanystropheus cf. longobardicus.
TANYSTROPHEUS LONCOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
43
Fig. 35 — Tanystropheus cf. longobardicus , MSNM V 3730, right pes in piantar view. I-V — metatarsals. Scale bar 10 mm. Preparation:
Giuseppina Damiano. Photo: Luciano Spezia.
44
STEFANIA NOSOTTI
DISCUSSION
Discussion and interpretation of thè skull anatomy of
Tanystropheus : a new reconstruction
In spite of thè large number of specimens of T. longo-
bardicus recovered during excavations in thè Besano
Formation ( =Grenzbitumenzone in thè Swiss geologi¬
ca! literature) thè strong compression of these fossils
together with thè frequent disarticulation of various
elements have been limiting factors in our interpreta¬
tion. This is especially true for thè skull. Skulls of T.
longobardicus in thè PIMUZ Collections that are more
or less complete are those of thè large-sized specimens
PIMUZ T 2790 (Wild, 1974: pi. 14) and PIMUZ T
2819 (Wild, 1974: figs. 5-6, pls. 15-16), and that of thè
small-sized specimen PIMUZ T 2791 (Wild, 1974: figs.
3-4; Fig. 38 in this paper). The latter was fìrst briefly
described by Peyer (1931). Wild (1974) gave a more
complete description, mainly based on X-radiographic
plates (Wild, 1974: fig. 3). However, this specimen is
severely crushed, and its interpretation very difficult.
Fig. 36 - Tanystropheus longobardicus , reconstruction of thè skull in thè small-sized specimens. See discussion on pp. 44-62.
Drawing: Massimo Demma.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
45
Another quite complete small-sized skull, PIMUZ T 3901
(Fig. 39), is somewhat better preserved. This specimen,
however, was referred by Wild (1980a) to a separate spe-
cies, T. meridensis (see “Introduction” for discussion).
The new specimens from Besano described here, par-
ticularly thè remarkably well preserved MSNM BES SC
1018, offer valuable new evidence on thè morphology of
thè skull in Tanystropheus .
In providing this new reconstruction of thè skull (Figs.
36-37), three points should be considered.
Firstly, thè reconstruction of thè skull of Tanystro¬
pheus presented here is only applicable to small-sized
representatives of T. longobardicus (or, altematively, to
thè representatives of a small-sized species of Tanystroph¬
eus , see p. 8) and also to T. meridensis. Differences in thè
shape and/or size of skull elements between small- and
large-sized PIMUZ specimens were observed but are not
discussed in any detail.
Secondly, thè ventral and occipital views of thè skull
were not reconstructed in detail. The elements of thè
dermal palate and of thè neurocranium are too poorly pre¬
served and/or exposed in thè new specimens, so they add
nothing further to thè previous descriptions. The extent of
thè dermal palate and thè occiput in thè model is consist-
ent with thè reconstruction of Wild (1974: fig. 8).
Thirdly, in thè words of Underwood (1970: 2): when
looking at thè reconstruction, “I hope that readers will
accept thè more extended flights of speculation as intended
to provoke discussion rather than to state positions firmly
held”.
The drawings in Figs. 36 and 37 were developed from
a clay model, that was used to test thè proportions and
thè nature of thè contacts between adjacent elements of
thè skull in thè three dimensions. The final reconstruction
resulted from a complex procedure, in which different
actions were performed simultaneously, each affecting
each other, so that slight adjustments were continu-
ously necessary. As a first step, an unfinished model was
made approaching thè generai proportions of thè skull in
MSNM BES SC 1018 and in PIMUZ T 2484, as recon¬
structed by Wild (1974: figs. 7 a, 8). Indeed, PIMUZ T
2484 shows very similar proportions of thè skull bones
to those of MSNM BES SC 1018. Precise paper tem-
plates of thè bones were then made from MSNM BES
SC 1018 (Figs. 10-13) and PIMUZ T 2484 (Figs. 40, 43,
48, 50-52), and applied to thè rough model, which was
consequently modified. The three-dimensional propor¬
tions of thè individuai skull elements and their respec-
tive contacts were finally adjusted working directly on
thè model, paying attention to how each change affected
thè overall configuration of thè skull. The squamosal was
reconstructed based on MSNM BES SC 265 (Figs. 2-3)
and PIMUZ T 279 1 (Fig. 38).
Premaxilla and Maxilla
The premaxilla and maxilla of MSNM BES SC 1018
are very similar in proportions and shape to thè same
elements in PIMUZ T 2484 (Wild, 1974: figs. 2, 82, 83,
pi. 18; Fig. 43 in this paper), in PIMUZ T 2791 (Wild,
1974: fig. 4; Fig. 38 in this paper) and in T. meridensis
Fig. 37 - Tanystropheus longobardicus, three-dimensional reconstruction of thè skull in thè small-sized specimens. See discussion on
pp. 44-62. Watercolor: Massimo Demma.
L
46
STEFANIA NOSOTTI
Fig. 38 — Tanvstrophens longobardicus , PIMUZ T 2791, skull. Scale bar 10 mm. Photo: Heinz Lanz, PIMUZ.
(Wild, 1980a: fig. 1; Fig. 39 in this paper). There are
some distinctive features common to all these speci-
mens but not present in thè large-sized individuals. The
premaxilla forms a long and slender maxillary process,
and a short, yet distinct nasal process. The maxilla is
triangular in shape anteriorly - without a premaxillary
process ( sensu Wild, 1974: 9, fig. 82) - and forms a very
well developed dorsal process.
The nature of thè suturai contact between thè premax¬
illa and thè maxilla is quite clear, and invariably thè same
in all thè skulls of thè small-sized specimens. As can be
seen in PIMUZ T 2484 (Fig. 43), thè antero-lateral sur-
face of thè maxilla has a deep furrow, matching thè maxil¬
lary process of thè premaxilla. T. meridensis (Fig. 39), in
which thè contact between thè premaxilla and maxilla can
be seen both laterally (left side of thè skull) and medially
(right side of thè skull), confirms that thè maxillary proc¬
ess of thè premaxilla overlaps thè maxilla laterally, fitting
into thè maxillary furrow. According to Wild (1974: 9; see
also reconstructions in figs. 7 a, 8, 10 a) thè dorsal proc¬
ess of thè maxilla were overlapped by thè nasal and thè
prefrontal. However, thè nature of thè contact between
thè three bones remains not completely clear, and cannot
be seen in either MSNM BES SC 1018 or in any PIMUZ
specimen. My hypothesis is that thè dorsal process of thè
maxilla was not concealed by thè nasal and prefrontal (see
“Nasal” and “Prefrontal”).
Nasal
In thè majority of known T. longobardicus speci¬
mens thè nasals are not preserved. A fragmentary, iso-
lated nasal is preserved in thè large specimen PIMUZ
T 2818 (Wild, 1974: pi. 12), and both nasals are com¬
pletely preserved, albeit isolated, in thè small PIMUZ
T 2484 (Fig. 40). Finally, in T. meridensis thè nasals
are preserved in situ but severely crushed (Fig. 39).
According to Wild (1974: 9), a loose junction of thè
nasals with thè frontals would account for thè nasals
being invariably present as isolated elements (or com¬
pletely missing). It seems likely that thè nasals had
very loose contacts with all thè neighbouring bones.
Consequently, thè nature of these contacts is very dif-
ficult to reconstruct.
According to Wild’s hypothesis (1974: fig. 8) thè
nasals overlapped thè dorsal process of thè maxillae, and
were overlapped by thè prefrontals and thè frontals. In
thè known Tanystropheus specimens, however, there is
no evidence for thè nasal overlapping thè dorsal process
of thè maxilla. The latter is fully exposed both in PIMUZ
T 2791 (Fig. 38) and in MSNM BES SC 1018. In these
specimens no nasal is preserved. However, I assume that
thè nasals in T. meridensis are preserved in situ (Fig.
39), crushed medially against thè dorsal process of thè
maxilla which is also fully exposed (contra Wild, 1980a:
5, I clearly identified a well developed dorsal process of
thè maxilla in T. meridensis ). This condition ot preserva-
tion suggests that thè nasal was positioned mediai to thè
dorsal process of thè maxilla, rather than overlapping it.
It is possible that thè lateral margin of thè nasal and thè
dorsal margin of thè maxilla established a syndesmotic,
unossified contact. It is difficult to ascertain whether thè
nasals were overlapped by thè prefrontals and thè fron¬
tals, but I assume that such was thè case (see “Frontal”
and “Prefrontal”).
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRET ATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
47
STEFANIA NOSOTTI
48
Fig. 40 - Tanystropheus longobardicus, PIMUZ T 2484, nasals. Scale
bar 1 5 mm. Photo: Heinz Lanz, PIMUZ.
The generai shape and proportions of thè nasals
were drawn based on those of PIMUZ T 2484 (Fig. 40)
but slightly modified to fìt them into thè model and to
obtain elongate external nares. Elongation of thè exter-
nal nares was inferred from T. meridensis (Fig. 39), in
which thè entire length of thè maxillary process of thè
premaxilla forms thè latero-ventral margin of thè exter¬
nal naris.
Frontal
In MSNM BES SC 1018 thè frontals are paired bones,
as they are in thè small-sized PIMUZ specimens (Wild,
1974: 10), in which thè suture between thè two elements
is partly identifiable.
In all specimens of T. longobardicus thè frontals
display large lateral axe-shaped flanges {die Orbital-
lamellae or orbitai laminae sensu Wild, 1974: 10).
Wild (1974: 10) stated that these flanges were flattened
because of compression but were originally slanting
latero-ventrally, (Wild, 1974: figs. 7 a, 8). By contrast,
I think that thè condition of preservation reflects thè
originai shape of thè frontals. In all thè PIMUZ speci¬
mens, isolated frontals are invariably preserved with
horizontal lateral flanges (Figs. 41-43). In thè complete
skulls, preserved in lateral or dorso-lateral view, there
is no evidence for a vertical orientation of thè flanges.
Fig. 41 — Tanystropheus longobardicus , PIMUZ T 2482, fronto-parietal piate in ventral view. The arrow points to thè fronto-parietal
suture. Scale bar 6 mm. Photo: Nicholas C. Fraser.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETAT10NS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
49
Fig. 42 - Tanystropheus longobardicus, PIMUZ T 2787, fronto-parietal piate in ventral view. Scale bar 10 mm. Photo: Nicholas
C. Fraser.
In T. meridensis (Fig. 39) thè two frontals rotateci as a
single unit and they clearly lie on one piane, with thè
dorsal surface exposed. The same is probably trae for
PIMUZ T 2791 (Wild, 1974: figs. 3-4; Fig. 38 in this
paper), although this specimen is difficult to interpret
because of extensive crushing. In MSNM BES SC 1018
(Figs. 10-12) there is no evidence that thè lateral flanges
were oriented latero-ventrally. The left frontal was
raised upwards, exposing thè entire ventral surface of
thè lateral flange, rather than extending into thè orbit as
it would have been expected if thè lateral flange formed
a mediai wall to it. In MSNM BES SC 265 (Figs. 2-3),
which is preserved in ventral view, thè lateral flanges of
thè frontals are seen as bony laminae roofing thè orbits.
The horizontal orientation of thè lateral flanges of thè
frontals in thè model (Figs. 36-37) is consistent with thè
generai proportions of thè skull.
The frontal piate of MSNM BES SC 1018 was
reconstructed duplicating thè left frontal (exposed in
ventral view). As a result, thè frontal piate in ventral
view (Fig. 44) closely resembles that of PIMUZ T 2787
(Fig. 42). Paired sinuous ridges characterize thè ventral
surface of thè frontals (Figs. 41-42, 44). These ridges
prosecute posteriorly on thè ventral surface of thè pari-
etals (see “Parietal”). Contra Wild (1974: 10). I do not
consider these ridges to mirrar structures on thè dorsal
surface of thè frontal, in particular thè supraorbital
ridges or die Supraorbitalkanten sensu Wild (1974:
10, fig. 2). According to Wild, thè supraorbital ridges
on thè dorsal surface of thè frontals formed thè dorsal
margin of thè orbits and of thè vertical orbitai lami¬
nae and they were impressed upon thè ventral surface
through compression. It is not clear what constitutes
thè cristae cranii frontalis considered by Wild (1974:
10) a feature of thè ventral surface of thè frontals.
Unequivocal evidence that thè sinuous ridges repre-
sent reai features of thè ventral surface of thè frontal
piate is provided by thè correspondence of thè skeletal
anatomy to that of thè soft parts. By comparison with
casts of thè endocranial cavity in extinct and extant
reptiles (Hopson, 1979), it is clear that thè shape of
thè sinuous ridges on thè frontal piate corresponds to
thè outline of thè forebrain (see also Wild, 1974: 10).
This is further confirmed by thè presence on thè ventral
surface of thè anterior process of thè frontal piate (see
below) of thè ovai impressions of thè olfactory bulbs
50
STEFANIA NOSOTTI
Fig. 43 - Tanystropheus longobardicus, PIMUZ T 2484, fronto-parietal piate in dorsal view. The arrows point to articular facets, see
discussion on p. 51. Scale bar 10 mm. Photo: Nicholas C. Fraser.
(Figs. 41-42). Hopson (1979: figs. 1-2) compared
endocasts made from thè skull of small and medium-
sized specimens of thè extant Caiman crocodilus and
observed that changes in endocast proportions reflect
changes in forebrain shape during ontogeny. In particu-
lar, in smaller individuals thè cerebral cast is propor-
tionally wider and in larger individuals it is narrower.
Interestingly, thè shape of thè sinuous ridges on thè
ventral surface of thè frontal piate in thè small PIMUZ
T 2482 is slightly different from thè larger PIMUZ
T 2787. This might be due to individuai variation or
alternatively be related to changes in forebrain shape.
In thè frontal piate of T. meridensis (Fig. 39), exposed
in dorsal view, Wild (1980a) described a thick supraor-
bital ridge continuous with thè orbitai margin of thè pre-
frontal. I interpret this ridge as thè mediai contact between
thè two frontals. The mediai relief on thè frontal piate of
T. meridensis is at least partly due to compression of thè
two frontals against thè other. However, thè right frontal
of MSNM BES SC 1018, with a strongly thickened and
sculptured mediai margin, suggests that there was a raised
area where thè two frontals met.
A tripartite anterior process (nasal process sensi i Wild,
1974: 10) formed by thè two frontals such as seen in
PIMUZ T 2484 (Fig. 43) is apparently not conspicuous in
MSNM BES SC 1018. The shape of this process results
from thè contact of thè anterior ends of thè frontals. As
described in MSNM BES SC 1018 (p. 21, Fig. 12), thè
anterior end of thè main body of thè frontal is bifurcate,
forming two lobes. In this specimen thè lateral lobe is
seen lateral (dorsal, as preserved) and thè mediai lobe
is seen mediai (ventral, as preserved) to thè interlocking
prefrontal. The mediai lobe in MSNM BES SC 1018 is
apparently very short, so that if thè anterior end of thè
left frontal is duplicated, a very short anterior process
of thè frontal piate results. I emphasize, however, that
an elongate triangular fragment of bone is preserved in
MSNM BES SC 1018 anterior to thè mediai lobe (Fig.
11), and this might be interpreted as an anterior continu-
ation of thè lobe itself, which as a consequence would be
longer. Considering also that a tripartite, well developed
anterior process of thè frontal piate is invariably present
in all PIMUZ specimens (Figs. 41-43), I reconstructed it
in thè model.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
51
The dorsal surface of thè anterior process can be seen
in PIMUZ T 2484 (Fig. 43). It shows paired articular
facets on each side, one between thè mediai and lateral
lobes, thè other in thè notch between thè anterior process
and thè lateral flange of thè frontal. Wild (1974: 10, fig.
2) thought that thè fronto-parietal piate of PIMUZ T 2484
was preserved in ventral view, and related thè presence
of thè facets on thè anterior process of thè frontal piate
to a contact with thè nasals and prefrontals. According to
Wild (1974: 9-10), both these elements were overlapped
by thè frontals. He assumed that thè anterior process of
thè frontal piate “fit[ted] in between thè nasals on thè
dorsal side”, and that it was almost completely covered
by thè prefrontals on thè ventral side. However, Wild did
not state precisely which elements were received in thè
articular facets on thè anterior process, and concluded that
thè nature of thè contacts between thè frontals, nasals and
prefrontals was very complex.
The mediai articular facets on thè dorsal side of thè
anterior process of thè frontal piate in PIMUZ T 2484
might be for thè receptions of thè nasals. However, thè
mediai margin of thè nasals of PIMUZ T 2484 (Fig. 40)
is not notched posteriorly. This indicates that if thè articu-
lated nasals had overlapped thè anterior process of thè
frontal piate they would have completely concealed it. By
contrast, evidence from PIMUZ T 2484 is that thè median
part of thè anterior process between thè two articular
facets was exposed on thè dorsal side of thè skull roof.
Consequently, I assume that thè nasals were overlapped
by thè anterior process of thè frontal piate. However, thè
nature of thè contact between thè nasals and thè frontals
remains unclear.
In MSNM BES SC 1018, thè prefrontal and thè fron¬
tal are stili fully articulated (p. 24) in an interlocking
suture, exposed in ventral view. The frontal process of
thè prefrontal is received into a notch between thè mediai
and lateral lobes of thè anterior process of thè frontal. In
dorsal view, this notch corresponds to thè margin of thè
articular facet between thè two lobes. I assume that thè
frontal process of thè prefrontal was directed medially
and contacted thè anterior process of thè frontal in an
oblique suture. Dorsally, thè prefrontal overlapped thè
mediai facet of thè anterior process, ventrally, thè margin
of thè prefrontal met thè margin of thè mediai facet.
The lobe of thè prefrontal (p. 24, Figs. 12, 37) appar-
ently does not reach thè facet on thè notch between
thè anterior process and thè lateral flange of thè fron¬
tal. Moreover, thè margin of thè lobe is thickened and
rounded, suggesting that it was probably free. At thè same
time, if thè lobe of thè prefrontal did not reach thè lateral
facet on thè anterior process of thè frontal piate, thè latter
facet remains unexplained.
Based on thè well preserved right parietal of MSNM
BES SC 1018, thè frontals fitted posteriorly in a V-shaped
embayment formed by thè parietals (Fig. 44). As a result,
thè parietal foramen is partly delimited by thè frontals. A
similar morphology is seen in PIMUZ T 2482 (Fig. 4 1 ). By
contrast, in PIMUZ T 2484 (Fig. 43), preserved in dorsal
view, thè frontals contact thè parietals in an unequivocal
interdigitating suture which is positioned anterior to thè
parietal foramen. These differences in thè fronto-parietal
contact are probably due to individuai variation.
The posterior contact of thè frontals with thè postfron-
tals is discussed under thè heading “Postfrontal”.
Parietal
The parietals of MSNM BES SC 1018 and MSNM
BES SC 265 conform to Wild’s statement (1974: 11)
that thè parietal piate is formed by two separate ele¬
ments. They are completely fused in thè large-sized
specimens in thè PIMUZ Collections, while a suture
between thè two elements is stili partly identifiable in
thè small-sized specimens.
The parietals were reconstructed in ventral view
(Fig. 44), largely based on MSNM BES SC 1018
(right parietal, Figs. 10-12). The reconstructed pari¬
etals are very similar to those of PIMUZ T 2482 (Fig.
41; see also Wild, 1974: fig. 2, where, contra Wild,
thè parietals drawn in ventral view are clearly those of
PIMUZ T 2482, not of PIMUZ T 2484). The assump-
tion that thè parietals of PIMUZ T 2482 are preserved
in ventral view is consistent with thè shape of thè
frontals in thè same view, as interpreted above. As for
thè frontals, I consider that thè condition of preserva-
tion of thè parietals reflects their originai morphol¬
ogy, and that thè relief on thè ventral surface of thè
parietals is not a direct result of compression causing
thè relief on thè dorsal surface to be impressed upon
it. The morphology of thè ventral surface of thè pari¬
etals conceivably mirrors that of thè soft parts in thè
cerebellar region of thè brain (see also “Frontal” and
Wild, 1974: 11).
mio
Fig. 44 - Tanystropheus longobardicus, reconstruction of thè fronto-
parietal piate in ventral view based on MSNM BES SC 1018. Water-
color: Massimo Demma.
r
52
STEFANIA NOSOTTI
In thè model, thè contact between thè parietals and
thè frontals was reconstructed based on MSNM BES SC
1018 (see “Frontal”), while that between thè parietals
and thè postfrontals (see “Postfrontal”) was inferred
from PIMUZ T 2484 (Fig. 43). The lateral supratempo-
ral processes were reconstructed on thè basis of MSNM
BES SC 265 and thè PIMUZ specimens. The supratem-
poral processes overlap thè squamosals and possibly
thè supratemporals (see “Supratemporal”), as can be
inferred from thè presence of a facet on their ventral
surface in MSNM BES SC 265 (Figs. 2-3) and PIMUZ
T 2482 (Fig. 41), and from thè articulated squamosal in
thè large PIMUZ T 2819 (Wild, 1974, figs. 5-6). Finally,
in thè parietals in dorsal view I assumed thè presence
of lateral obliquely slanting flanges {die TemporalfliÀgel
or parietal laminae sensu Wild, 1974: 11) but there is
no clear evidence for them either in thè new specimens
or in thè small-sized PIMUZ specimens. They are only
unequivocally present in thè large PIMUZ T 28 1 9 (Wild,
1974: figs. 5-6).
The ratio between thè length of thè parietals relative
to thè overall length of thè fronto-parietal piate result-
ing from thè reconstruction in MSNM BES SC 1018 is
congruent with that observed in PIMUZ T 2484 (Fig.
43). In thè larger PIMUZ T 2482 (Fig. 41) and PIMUZ T
2787 (Fig. 42) thè ratio is progressively higher. The rela¬
tive length of thè frontal and thè parietal changes, with
thè parietal increasing and thè frontal decreasing with
increasing overall size.
Supratemporal
Wild (1974: 12) claimed thè presence of supratempo¬
rals in T. longobardicus on thè basis of an element inter-
preted as a supratemporal preserved in situ in thè large-
sized PIMUZ T 2819. Paired rod-shaped small bones
might be interpreted as supratemporals in MSNM BES
SC 265 (p. 12, Figs. 2-3). For thè present thè occurrence
of supratemporals in T. longobardicus is equivocai and
it is therefore impossible to describe thè nature of their
contacts with thè neighbouring bones.
Prefrontal
The prefrontal of MSNM BES SC 1018 is very similar
to thè prefrontal of T. meridensis (Fig. 39) but different
from thè prefrontals as described and figured by Wild
(1974: 10, figs. 8-9) in thè specimens of T. longobardicus
in thè PIMUZ Collections.
In Wild’s reconstructions (1974: figs. 8-9), thè pre¬
frontals extend in a antero-posterior direction and only
form thè antero-dorsal margin of thè orbits. Further ven¬
tral ly, thè anterior margins of thè orbits are formed by thè
lacrimals. The prefrontals overlap both thè dorsal process
of thè maxillae and thè nasals with a process (anterior
process) extending anteriorly far beyond thè fronto-
nasal contact. Medially, they contact thè frontal (frontal
process), extending posteriorly beyond thè fronto-nasal
contact. The “orbitai lamina” of thè prefrontal is con-
tinuous with thè “orbitai lamina” of thè frontal. Wild also
maintained that thè prefrontals reached thè palatine with
a palatine process.
Among thè small-sized PIMUZ specimens, Wild
(1974: 10) identifìed a prefrontal preserved in situ in
PIMUZ T 2485 and an isolated prefrontal in PIMUZ T
2795. It is on thè latter that he based thè drawing in fig. 2
and thè reconstruction in fig. 8. However, in Wild’s draw¬
ing of PIMUZ T 2485 (Wild, 1974: pi. 8) a prefrontal is
not identifiable and in thè radiograph of PIMUZ T 2795
(Wild, 1974: pi. 6) it is diffìcult to understand which ele¬
ment Wild was referring to as thè prefrontal.
In MSNM BES SC 1018 (Figs. 10-12) and in T.
meridensis (Fig. 39) thè prefrontal mostly extends in
a dorso-ventraf direction and forms almost thè entire
anterior margin of thè orbit. Based on MSNM BES
SC 265 (p. 12, Figs. 2-3), I confimi that thè prefrontal
reached thè palatine ventrally. In MSNM BES SC 1018
and in T. meridensis thè prefrontal is located posterior
to thè dorsal process of thè maxilla, and not overlapping
it. Based on thè interpretation of thè contact between
thè frontal and thè prefrontal (see “Frontal”), thè latter
element had to overlap thè nasal in order to reach thè
anterior process of thè frontal piate. The reconstruction
presented here differs from Wild’s (1974: figs. 8-9) in
that thè prefrontal does not extend far anteriorly beyond
thè fronto-nasal contact. Finally, thè prefrontal is recon¬
structed in contact with thè anterior process of thè fron¬
tal piate only, positioned mainly anterior to thè latter,
not lateral to it. In thè new reconstruction thè “orbitai
laminae” of thè frontal are horizontally, not vertically
positioned as was assumed by Wild. Consequently, thè
orbitai surface of thè prefrontal contacts thè ventral sur¬
face of thè frontal more medially, and there is no conti-
nuity between an “orbitai lamina” of thè prefrontal and
an “orbitai lamina” of thè frontal.
Among thè large-sized PIMUZ specimens, Wild
(1974: 10) identifìed thè prefrontals in PIMUZ T 2787.
According to Wild, thè right prefrontal is isolated but
complete (Wild, 1974: fig. 1), and it is very similar in
shape to thè prefrontal of thè small-sized specimens
(Wild, 1974: fig. 2). Wild also maintained that thè left
prefrontal of PIMUZ T 2787 (Fig. 42) is preserved in
situ, and that it stili contacts thè maxilla and thè frontal.
However, he stated that it was not possible to unequivo¬
cally identify thè margins of thè three bones. Indeed, I
could not identify any contact between thè maxilla and
thè presumed prefrontal. Wild assumed that thè prefron¬
tal widely overlapped thè ventral surface of thè anterior
process of thè frontal but thè presumed prefrontal of
PIMUZ T 2787 does not. An alternative interpretation
might be that thè “prefrontal” in PIMUZ T 2787 is in fact
a nasal but also in this case thè nature of thè contact with
thè frontal is not clear.
Lacrimai
A lacrimai was only reported by Wild (1974: 10, figs.
1, 2) for PIMUZ T 2795 and PIMUZ T 2787, and in both
specimens it is an isolated bone. The lacrimai of PIMUZ
T 2795 was figured by Wild (1974: fig. 2) as a triangular
bone with no surface detail. The shape of thè lacrimai
of PIMUZ T 2787 as figured by Wild (1974: fig. 1) is
comparable in shape to that of MSNM BES SC 1018,
although thè lacrimai foramen is more posteriorly located
in PIMUZ T 2787. The lacrimai of MSNM BES SC
1018 documents for thè first time a lacrimai of T. longo¬
bardicus preserved in situ, confirming Wild’s reconstruc¬
tion ( 1 974: fig. 8 b). A lacrimai of approximately thè same
shape, size and position as that in MSNM BES SC 1018
was identifìed by Wild (1980a: fig. 1) in thè skull ot T.
meridensis (Fig. 39).
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRET ATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
53
Postfrontal
The new specimens provide very limited information
on thè morphology of thè postfrontal, and no postfron-
tals are preserved in thè majority of PIMUZ specimens.
However, in PIMUZ T 2484 postfrontals are preserved
stili sutured to thè frontals (Fig. 43; see also Wild, 1974:
fig. 2). This specimen best clarifies thè position, shape
and contacts of thè postfrontals in dorsal view. The post¬
frontals are subtriangular elements anteriorly contacting
thè horizontal flanges of thè frontals, and medially thè
parietals (Fig. 43). The exact same configuration can be
seen on thè radiograph of thè slightly larger PIMUZ T
2482 (Fig. 45). An articulated postfrontal is probably also
preserved in thè small PIMUZ T 2791 (Wild, 1974: 18,
figs. 3-4; Fig. 38 in this paper) but this skull is severely
crushed and thè information it provides is limited. Finally,
a postfrontal is probably preserved in T. meridensis (Wild,
1980a; Fig. 39 in this paper).
Fig. 45 - Tanystropheus longobardicus , PIMUZ T 2482, fronto-pari-
etal piate, radiograph. The arrow points to thè postfrontal. Courtesy
PIMUZ.
In thè large PIMUZ T 28 1 9 (Wild, 1 974: figs. 5-6), thè
lefìt postfrontal appears to be at least partially preserved
but it provides little detail. Wild (1974) claimed thè pres-
ence of a postfrontal in PIMUZ T 2787 (Fig. 42), but
there is no evidence of a suture between thè frontal and
thè postfrontal as shown in Wild’s fig. 1. Evidence from
other specimens indicates that thè presumed postfrontals
in PIMUZ T 2787 represent thè posterior part of thè hori¬
zontal flanges of thè frontals. In thè large MSNM V 3663
(p. 38, Fig. 31) thè right postfrontal is preserved, sutured
to thè skull roof in a manner similar to PIMUZ T 28 1 9.
The postfrontal is therefore a very short element pro-
jecting laterally and ventrally, and sutured to thè frontal
and parietal. Its shape is similar in small- and large-sized
specimens. However, thè nature of its contact with thè
postorbitai is not completely clear, due to thè difficulties
encountered in thè interpretation of thè temporal region of
thè skull (see below).
Squamosal and Postorbitai
Wild (1974: 11) described thè squamosal as a sickle-
shaped element with a wide anterior postorbitai process
overlapping thè postorbitai, a pointed posterior parietal
or supratemporal process overlapped by thè supratem-
poral process of thè parietal, and a quadrate process
overlapping thè cephalic condyle of thè quadrate. Wild’s
description was evidently based on thè small PIMUZ T
2791 (Wild, 1974: figs. 2-4; Fig. 38 in this paper), thè
larger PIMUZ T 2787 (Wild, 1974: fig. 1, pi. 9; Fig. 42
in this paper) and thè large PIMUZ T 2819 (Wild, 1974:
figs. 5-6).
The generai shape of thè left squamosal in MSNM
BES SC 265 (p. 12) is very similar to that of thè squa¬
mosal in Wild’s (1974) fig. 2 (this element is not from
PIMUZ T 2484, as stated by Wild but clearly corresponds
to thè isolated right squamosal of PIMUZ T 2791, see Fig.
38 in this paper). I suggest that this shape represents reai
features of thè bone, and that thè squamosal originally
had a complex three-dimensional shape (Fig. 46) which
cannot be seen in thè fossils because thè bone is crushed
on thè slab. The orientation of thè posterior and anterior
rami of thè squamosal - and of its postorbitai process - in
thè three dimensions shown in Fig. 46 might well pro¬
duce thè shape of thè compressed squamosal described
for MSNM BES SC 265 (p. 12). As thè left squamosal of
MSNM BES SC 265 is preserved in its originai antero-
posterior orientation, there is unequivocal evidence for
thè pointed process of thè squamosal being thè postorbitai
process (contra Wild, 1974: 11, fig. 2).
Wild (1974: 1 1, fig. 2) interpreted thè postorbitai as an
elongate, subtriangular and flattened bone. Its long and
pointed ventral jugal process overlapped thè jugal, and
thè expanded dorsal end formed an anterior supraorbital
process (orbitai process sensu Wild, 1974), overlap¬
ping thè postfrontal, and a posterior squamosal process,
interlocking with thè squamosal. Among thè small-sized
specimens. Wild identified isolated postorbitals in
PIMUZ T 2484 (Wild, 1974) and T. meridensis (Wild,
1980a). While thè element interpreted by Wild (1980a:
fig. 1 ; Fig. 39 in this paper) as a postorbitai in T. meriden¬
sis is probably a quadrate, it is not clear which element of
PIMUZ T 2484 was identified by Wild as thè postorbital.
In thè caption of fig. 1 Wild ( 1 974) stated that thè isolated
postorbital illustrated was from PIMUZ T 2484 but he
did not indicate which bone he identified as thè postor¬
bital in thè “exploded” skeleton (Wild, 1974: pi. 18). It is
possible that it is thè element preserved to thè right side
of thè fronto-parietal piate (labelled “po?” in Fig. 43),
which is here considered as thè right postorbital. In thè
same specimen there are remains of a bone overlapped
by thè left prefrontal which conceivably represent thè left
postorbital (see Fig. 43 in this paper and Wild, 1974: pi.
18, where thè remains are labelled “postfrontal”). In thè
large-sized specimens in thè PIMUZ Collections, Wild
reported a postorbital for PIMUZ T 2819 (Wild, 1974:
fig. 6). This fragmentary element, however, is in all prob-
ability a squamosal (Kuhn-Schnyder, 1967: fig. 3). Wild
(1974) probably based thè description of thè postorbital
54
STEFANIA NOSOTTI
Fig. 46 — Tanys troph e us longobardicus, reconstruction of thè left
squamosal in thè small-sized specimens (see discussion on p. 53).
A) Dorsal view. B) Ventral view. Drawing: Massimo Demma.
on thè larger-sized PIMUZ T 2787 (Wild, 1974: fìg. 1,
pi. 9), in particular on thè element labelled “po?" in
Fig. 47. Two additional elements preserved in PIMUZ
T 2787 (Fig. 42) were interpreted by Wild as thè squa-
mosals. The element labelled “po” in Fig. 42, however,
is similar in shape to thè putative postorbitai described
for MSNM BES SC 1018 (p. 24). Contra Wild (1974), I
suggest that it represents a postorbital. Its shape is difter-
ent from that of thè element labelled “sq” in Fig. 42. In
this latter element a laterally projecting posterior ramus
and an antero-laterally oriented anterior ramus - with a
tapering postorbital process bending medially - can be
clearly seen. Therefore, I concur with Wild (1974) that
this second element is indeed a squamosal. The preserved
part of thè right postorbital in MSNM V 3663 (Fig. 31)
is conceivably thè dorsal one, extending between thè
postfrontal and thè squamosal, while thè jugal process is
assumed to be truncated.
The contacts of thè squamosal and postorbital with thè
neighbouring bones are not completely clear.
Wild (1974: 11, 18) stated that both in thè small-
sized PIMUZ T 2791 (Wild, 1974: figs. 3-4) and in thè
large-sized PIMUZ T 2819 (Wild, 1974: figs. 5-6) there
were squamosals preserved in situ, and he maintained
that thè articular relationships between thè squamosal
and thè neighbouring bones, as indicated in thè recon-
structions presented in figs. 8 and 9 (Wild, 1974), were
derived from these specimens. Contra Wild, I suggest
that these elements only contact thè parietal. Assuming
Wild correctly identified thè left squamosal in PIMUZ T
2791 on thè stereo-radiographs (Wild, 1974: figs. 3-4),
this element is oriented in thè direction of thè ascend-
ing process of thè jugal, in a manner similar to thè left
squamosal in MSNM BES SC 265. It does not contact
thè postorbital antero-ventrally. In fact, a postorbital
as described by Wild cannot be identified in PIMUZ
T 2791. The element identified by Wild as thè right
squamosal in PIMUZ T 2819 is broken anteriorly, and
again a right postorbital was not described by Wild in
this specimen.
As in all thè available material of T. longobardicus
there are no articulated squamosals and postorbitals, thè
nature of thè contact between these two elements remains
unclear. However, thè element tentatively identified as
thè right postorbital in PIMUZ T 2484 (Fig. 43) exhibits
a posterior triangular and longitudinally elongate facet.
This might be a facet for thè reception of thè postorbital
process of thè squamosal.
Wild emphasized (1974: 11) that thè articular rela-
tionship between thè postorbital and thè postfrontal was
not clear. Assuming that thè fronto-parietal piate and thè
postfrontals in PIMUZ T 2484 (Fig. 43) were preserved in
ventral view. Wild conjectured that thè postorbital over-
lapped thè dorsal surface of thè postfrontal, because on
thè ventral surface of thè latter a facet for thè reception of
thè postorbital cannot be seen. Contra Wild, I assume that
thè fronto-parietal piate and thè postfrontals in PIMUZ
T 2484 are preserved in dorsal view. This would suggest
that thè postorbital was overlapped by thè postfrontal.
However, thè presumed left postorbital is very fragmen-
tary, and thè nature of thè contact between thè postfrontal
and thè postorbital cannot be stated precisely. Articulated
postfrontals and postorbitals are not preserved in other
PIMUZ specimens. Specimens MSNM BES SC 265 and
TANYSTROPHEUS LONGOBARD1CUS. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
55
Fig. 47 - Tanystropheus longobardicus , PIMUZ T 2787, presumed postorbitai (see discussion on p. 54). Scale bar 20 mm. Photo:
Nicholas C. Fraser.
MSNM BES SC 1018 add no information. Apparently,
only MSNM V 3663 preserves a postfrontal in articu-
lation with thè postorbitai (p. 38). According to Wild
(1974: 11), a loose junction of thè postorbitai with thè
neighbouring elements would account for thè postorbitai
being mostly preserved as an isolated element. In MSNM
V 3663, however, thè postfrontal and thè postorbitai seem
to be steadily sutured (Fig. 31).
No articulated postorbitals and jugals are preserved in
thè PIMUZ material and thè configuration of thè contact
between thè two elements was here inferred from MSNM
BES SC 1018 (p. 24).
The result of my interpretation of thè temporal region
of thè skull is shown in Figs. 36-37. The posterior ramus
of thè squamosal and thè posterior part of its anterior
ramus He on a piane slanting downwards. The anterior
part of thè anterior ramus of thè squamosal (postorbitai
process) is deflected ventrally and medially, where it con-
tacts thè posterior side of thè postorbital. The postorbitai
is represented as a triradiate element. Dorsally and ante-
riorly it contacts thè postfrontal. Ventrally, thè bifid jugal
process of thè postorbital contacts thè jugal. The anterior
ramus of thè jugal process fìts into thè elongate fossa on
thè dorsal process of thè jugal.
Comparison with thè skull of thè closely related Mac-
rocnemus (Rieppel & Gronowsky, 1981: fig. 1; Premru,
1991: fig. 3; Renesto & Avanzini, 2002: fig. 2 A) and thè
more distantly related Prolacerta (Modesto & Sues, 2004:
fig. 3) seems to lend further support for thè presence of
a triradiate postorbital and a short, triangular postfrontal
in Tanystropheus . Both in Macrocnemus and Prolacerta
thè two elements are in loose contact. Interestingly, in
thè uncatalogued Macrocnemus PIMUZ specimen “Alla
Cascina” (Rieppel & Gronowsky, 1981: fig. 1), thè pos¬
torbital process of thè squamosal contacts a long, hori-
zontal posterior process of thè postorbital along its entire
ventral margin.
Jugal
A jugal is preserved in lateral view both in MSNM
BES SC 265 and in MSNM BES SC 1018; in thè latter
specimen thè bone is particularly well preserved in situ.
The comparison with thè small- and large-sized
specimens in thè PIMUZ Collections revealed that a
much taller dorsal or postorbital process characterizes
thè small-sized specimens by comparison with large-
sized ones.
56
STEFANIA NOSOTTI
A feature that was not mentioned by Wild (1974)
because it cannot be seen in thè PIMUZ material is thè
presence of an elongate groove dose to thè orbitai margin
of thè dorsal process. This groove is posteriorly delimited
by a crest, which itself is positioned anterior to a shallow
elongate fossa.
Quadrate
The left quadrate of MSNM BES SC 1018 is badly
crushed but apparently similar in shape to thè quadrate
of T. meridensìs (Wild, 1980a: fig. 1; Fig. 39 in this
paper).
Wild (1980a) claimed that thè shape of thè quadrate
was one of thè diagnostic characters of thè new species
T. meridensìs. According to him, thè quadrate of T. meri-
densis would be taller, and more slender and concave
posteriorly than in T. longobardicus.
Among thè small-sized specimens of T. longobardicus
in thè PIMUZ Collections both quadrates are preserved
as isolated elements in PIMUZ T 2484 (Wild, 1974: fig.
2). One of thè two (left, Fig. 43), exposing its mediai side
according to Wild, is badly crushed. The other (right, Fig.
40), exposing its lateral side according to Wild, provides
more detail. As compared to thè quadrate of T. meriden-
sis, thè quadrate of T. longobardicus is apparently shorter
and wider only because of thè presence of a wide mediai
lamina. If thè ratio between thè height of thè quadrate
and thè length of thè maxilla is calculated, thè height
of thè quadrate is approximately 50% of thè length of
thè maxilla in T. meridensìs , and approximately 40% in
PIMUZ T 2484. In Wild’s reconstruction (1974: fig. 8)
based on PIMUZ T 2484, thè height of thè quadrate is
approximately 45% of thè length of thè maxilla. Fitting
thè lower jaw into thè model it became apparent that a
height of thè quadrate of approximately 47% of thè length
of thè maxilla was required for thè quadrate to bridge thè
distance between thè squamosal and thè articular fossa
of thè lower jaw (assuming that thè lower jaw was not
distorted). Significantly, thè height of thè quadrate so
obtained falls within thè range of 40-50% of thè length
of thè maxilla observed in thè PIMUZ specimens. As thè
left quadrate of MSNM BES SC 1018 is badly crushed
and fragmentary, it is difficult to evaluate thè presence of
a mediai lamina and its extent. The mediai lamina of thè
quadrate in T. meridensìs is apparently very short. How-
ever, part of thè mediai lamina might stili be enclosed in
thè matrix in both these specimens. Finally, thè quadrate
of PIMUZ T 2791 (Wild, 1974: fìgs. 3-4; Fig. 38 in this
paper), which is preserved in situ and exposed in lateral
view, exhibits thè same concave posterior margin as thè
quadrate of T. meridensìs.
In conclusion, thè quadrate of T. longobardicus prob-
ably had a shape similar to that described by Wild (1974)
for thè PIMUZ specimens, with an even wider mediai
lamina in thè large-sized specimens compared to thè
small-sized ones. There is no appreciable differences in
thè shape and height of thè quadrate in T. longobardicus
and T. meridensìs.
According to Wild (1974: 12, 25) thè quadrate of
Tanystropheus was streptostylyc. It was loosely articu-
lated with thè squamosal and, probably, thè supratempo-
ral, and had ligamentous connections with thè opisthotic
and thè pterygoid. This interpretation cannot be verified
in thè new specimens.
Epipterygoid
The epipterygoid is not preserved in MSNM BES SC
1018. According to Wild (1974: 14, fig. 1), it is preserved
only in thè larger specimens among thè material housed
at thè PIMUZ. An isolated epipterygoid is preserved in
MSNM BES SC 265 (p. 12, Figs. 2-3), and, as stated by
Wild, it is a rod-shaped element with expanded ends. As
thè contacts of this bone with other elements of thè skull
cannot be ascertained, it was not included in thè recon¬
struction.
Neurocranium
Only some elements of thè neurocranium are pre¬
served in thè new specimens. In particular, in MSNM
BES SC 265 thè basisphenoid-parasphenoid complex is
clearly identifìable, while thè other elements are badly
crushed, and their interpretation is very difficult (p. 12). In
MSNM BES SC 1018 I could only identify thè supraoc-
cipital (p. 24).
To assess whether thè extent of thè occiput recon-
structed in thè model was consistent with thè proportions
of other regions of thè skull I attempted a reconstruction
of thè supraoccipital-exoccipitals-opisthotics complex
in MSNM BES SC 1018 by direct comparison of its
supraoccipital with that preserved in PIMUZ T 2484
(Fig. 43). I then extrapolated thè proportions of thè exoc-
cipitals and opisthotic from their dimension in PIMUZ
T 2484.
The lateral end of thè reconstructed paroccipital proc-
esses of MSNM BES SC 1018 resulted very dose to thè
quadrate but did not quite reach it. Wild (1974: fig. 8)
carne to thè same conclusion. However, my reasoning
was based on a paper template of thè supraoccipital-exoc-
cipitals-opisthotics complex, which does not take into
account thè thickness of thè bone. Therefore a contact
of thè paroccipital process with thè quadrate cannot be
excluded.
Sclerotic ring
Wild (1974: 17) drew attention to thè presence in
thè PIMUZ material of small, thin bony plates with a
polygonal or rounded shape, and tentatively interpreted
them as sclerotic plates. In T. meridensìs (1980a: figs. 1-2)
he again identified ten thin roundish-oval bony elements
positioned within thè orbit as sclerotic plates.
MSNM BES SC 1018 confìrms thè presence of a
sclerotic ring in Tanystropheus. This specimen is thè only
one in all thè available material that shows isolated plates
with a definitive shape. In particular, four plates preserved
dorsal to thè skull indicate a subrectangular shape. Other
elements, mostly with quite irregular shape, are preserved
within thè orbit.
The proposed reconstruction of thè sclerotic ring
(Figs. 36-37) should be considered provisional. Firstly,
thè possible size of an eye was determined on thè size
of thè reconstructed orbit, followed by an estimate of
thè diameter of thè cornea. While arranging thè subrec¬
tangular sclerotic plates around thè cornea, I established
that thè plates should follow thè eye curvature, and that
18 plates were required to maintain thè contact of thè
plates orbitally (sensu Underwood, 1970). As a result,
thè plates partly overlapped corneally (sensu Under¬
wood, 1970). The reconstructed pattern of overlap is
arbitrary.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
57
Lower jaw
Within thè small-sized T. ìongobardicus speci-
mens in thè PIMUZ Collections, complete lower jaws
in situ are preserved in PIMUZ T 2791 (Wild, 1974:
figs. 3-4; Fig. 38 in this paper). In this specimen thè
right lower jaw overlaps thè left one to a large extent.
Since thè skull is also heavily crushed, PIMUZ T
2791 provides very little information for thè recon-
struction of thè lower jaw. The left mandibular ramus
is preserved in situ, exposed in lateral view, in thè
small T. meridensis (Wild, 1980a: fig. 3; Fig. 39 in
this paper). Wild (1980a: 7; Fig. 49 F in this paper)
maintained that thè configuration of thè different ele-
ments in thè lower jaw of T. meridensis was similar to
T. ìongobardicus.
In other “exploded” skulls of small-sized T. longo-
bardicus specimens in thè PIMUZ Collections, more or
less complete lower jaws are preserved as isolated ele-
ments. Both dentaries, post-dentary parts, and isolated
bones of thè lower jaws are preserved in PIMUZ T 2484
(Figs. 48, 50-52). On thè basis of this specimen Wild
(1974: 40, figs. 15-16 a) reconstructed thè lower jaw in
lateral view for thè small-sized T. ìongobardicus speci¬
mens. Specimen PIMUZ T 2482 (Wild, 1974: pi. 5)
preserves thè complete left lower jaw in mediai view
(Wild, 1974: fig. 27 b), as well as part of thè dentary of
thè right lower jaw in lateral view. Indeed, PIMUZ T
2482 provides thè most detail of thè mediai surface of
thè lower jaw within thè PIMUZ material but it is not
exhaustively informative. The fragmentary mandibular
ramus of PIMUZ T 2779 (Peyer, 1931: fig. 3 in pi. 13,
text-fìg. 14) is too poorly preserved and adds no sub-
stantial information. Consequently, Wild reconstructed
thè lower jaw in mediai view for thè small-sized T.
ìongobardicus specimens only tentatively (Wild, 1974:
fig. 16 b), based on thè configuration of thè bones on
thè lateral side of thè lower jaw and on isolated man¬
dibular elements (Wild, 1 974: 40, fig. 1 5) from PIMUZ
T 2484 (coronoid and prearticular) and PIMUZ T 2795
(splenial).
Complete mandibular rami in lateral view are pre¬
served in thè large-sized specimens PIMUZ T 2787
(right lower jaw; Wild, 1974: pi. 9) and PIMUZ T
2819 (left lower jaw; Wild, 1974: figs. 5-6), while
in PIMUZ T 2793 (Wild, 1974: figs. 18-19) thè right
lower jaw is heavily damaged, and thè left one is
broken into two pieces. The lateral surface of thè
lower jaw of thè large-sized specimens of Tanystro-
Fig. 48 - Tanystropheus ìongobardicus , PIMUZ T 2484, lower jaws. Scale bar 10 mm. Photo: Rosi Roth, PIMUZ.
58
STEFANIA NOSOTTI
Fiu 49 - Lower jaws in small-sized specimens of Tanystropheus in lateral view, not to scale. A) Tanystropheus longobardicus, small-
sized specimens, reconstruction of Wild (after Wild, 1974: fig. 16 a). B) Tanystropheus longobardicus, reconstruction of thè author,
drawing: Massimo Demma. C) Tanystropheus longobardicus, MSNM BES SC 265, left lower jaw, photo: Luciano Spezia. D) Tanys¬
tropheus longobardicus, MSNM BES SC 265, right lower jaw, reflected horizontally for easier comparison, photo: Luciano Spezia^
E) Tanystropheus longobardicus, MSNM BES SC 1018, left lower jaw, photo: Luciano Spezia. F) Tanystropheus mendensis, P1MLZ
T 390 Ì, reconstruction of Wild (after Wild, 1980a: fig. 3). G) Tanystropheus meridensis, P1MUZ T 3901, left lower jaw, as interpreted
by thè author, photo: Heinz Lanz, PIMUZ. Pink lines: suture dentary-surangular. Green lines: presumed coronoid. Orange lines: suture
articular-prearticular. See discussion on pp. 57-61.
TANYSTROPHEUS LONGOBARDICUS. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
59
pheus was reconstructed by Wild (1974: 40, fig. 17)
on thè basis of PIMUZ T 2819, thè mediai surface
on thè basis of radiographs of thè right lower jaw of
PIMUZ T 2787, and of a small fragment of thè left
lower jaw of thè same specimen. Wild (1974: fig. 19)
also provided a reconstruction of thè lateral surface of
thè lower jaw of PIMUZ T 2793 but this is similar to
that of PIMUZ T 2819.
As reconstructed by Wild, thè nature of thè contacts
between thè elements of thè lower jaw, is similar in both
small- and large-sized specimens of Tanystropheus.
A new reconstruction of thè lower jaw in lateral view
is presented here, based on MSNM BES SC 1018 (Figs.
13, 49 E), MSNM BES SC 265 (Figs. 2-3, 49 C-D), and
PIMUZ T 2484 (Figs. 48-52), which is partly re-inter-
preted. The confìguration of thè mandibular elements on
thè mediai side of thè lower jaw remains not completely
clear but thè right lower jaw of MSNM BES SC 1018
(Fig. 53 B) provides new information on thè shape of thè
splenial and prearticular.
By comparison with Wild’s reconstruction (1974:
fig. 16; Fig. 49 A in this paper), my reconstruction
of thè lower jaw in lateral view (Fig. 49 B) differs on
three points. First, thè surangular overlaps thè dentary.
Second, thè angular is wider. Third, thè splenial con-
tributes to a larger extent to thè ventral margin of thè
lower jaw.
Wild (1974: 38, figs. 15-16) maintained that thè
dentary overlapped thè surangular, observing that
thè surangular was remarkably grooved anteriorly to
establish a strong contact with thè dentary. However,
on both mandibular rami of MSNM BES SC 265,
an oblique, dorsally convex suture is unequivocally
present (Fig. 49 C-D), which can be identified as thè
contact between thè dentary and thè surangular. The
same suture occurs in MSNM BES SC 1018 (left lower
jaw, Fig. 49 E), albeit less distinct, and in T. meriden-
sis (Fig. 49 G). In both isolated dentaries of PIMUZ
T 2484 (Fig. 48) an oblique, dorsally convex line sets
off a depressed area, suggesting that thè dentary was
overlapped by thè surangular. In thè post-dentary iso¬
lated part of thè left lower jaw of thè same specimen
(Figs. 49, 51 B), thè anterior part of thè surangular
displays a deepened surface dorsal to a sinuous line.
I interpret this line to represent thè suture between
thè dentary and surangular on thè lateral surface of
thè lower jaw. I assume that, if observed anteriorly,
thè surangular would be excavated by a deep groove
separating a mediai and a lateral bony laminae, which
are compressed one over thè other in thè fossil. The
mediai lamina would overlap thè mediai surface of thè
dentary, thè lateral would overlap thè lateral surface of
thè dentary. In other words, thè dentary would fit into
thè groove between thè two laminae. On thè mediai
surface of thè right lower jaw of MSNM BES SC 1018
(Fig. 53 B), however, thè nature of thè contact between
thè dentary and thè surangular is not clear. Perhaps, a
coronoid overlapped thè area where thè two bones met
(see below).
Joining paper templates of thè dentary and of post-
dentary part of thè jaw in PIMUZ T 2484, I observed
that where thè two elements met, on thè ventral margin
Fig. 50 - Tanystropheus longobardicus, PIMUZ T 2484, post-dentary part of thè left lower jaw in lateral view. Scale bar 5 mm. Photo:
Rosi Roth, PÌMUZ.
60
STEFANIA NOSOTTI
Fig. 51 - Tanvstropheus longobardicus, PIMUZ T 2484, lower jaws. The best preserved dentary is shown in association with thè best
preserved post-dentary part of thè lower jaw. A) Right dentary in lateral view, reflected honzontally. B) Post-dentary part ot thè left
dentary in lateral view. Photo: Rosi Roth, PIMUZ.
Fig. 52 — Tanystropheus longobardicus, PIMUZ T 2484, post-dentary
part of thè right lower jaw in medio-lateral view. Scale bar 5 mm.
Photo: Rosi Roth, PIMUZ.
of thè lower jaw there is a notch (Fig. 51). I assume
that this notch accommodated thè splenial, as stated by
Wild (1974: fig. 16). Indeed, Wild found poor evidence
for thè shape of thè splenial in thè PIMUZ material.
Based on thè right lower jaw of MSNM BES SC 1018
(Fig. 53 B), I conclude that this element extended along
thè ventral margin of thè lower jaw far more posteri-
orly than it was assumed by Wild (Figs. 49 A-B, 53).
Evidence from MSNM BES SC 1018 also indicates
that thè isolated element dose to thè posterior part
of thè right lower jaw in PIMUZ T 2484 (Wild, 1974:
fig. 15; Figs. 48, 52 in this paper), interpreted by Wild
as a prearticular, is more probably a fragment of thè
splenial.
The nature of thè contact between thè surangular
and thè angular in lateral view cannot be inferred from
thè new specimens but it is clear in PIMUZ T 2484. In
Fig. 53 - Lower jaw in mediai view, in small-sized specimens of Tanystropheus longobardicus. A) Reconstruction ot Wild
(modified after Wild, 1974: fig. 16 b). B) Right mandibular ramus of MSNM BES SC 1018 as interpreted by thè author, drawing:
Massimo Demma.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
61
both thè left (Figs. 50, 51 B) and thè right (Figs. 48,
52) isolated post-dentary parts of thè lower jaws of
this specimen I identifìed thè suture between thè two
elements as an oblique, dorsally convex line, which is
positioned more dorsally than assumed by Wild. In all
probability, he identifìed as a suture thè line delimiting
a wide ventral area characterized by a striking sculp-
tured surface for muscular attachment. This was prob-
ably for thè insertion of thè posteroventral lb (super-
fìcialis layer of thè external adductor) described by
Rieppel & Gronowsky (1981), and assumed by these
authors to be present also in Macrocnemus (Rieppel &
Gronowsky, 1981: fìg. 6 B), a taxon closely related to
Tanystropheus.
Posteriorly, as can be seen more clearly in thè iso¬
lated post-dentary part of thè left lower jaw of P1MUZ
T 2484 (Fig. 51 B), thè surangular meets thè articular
in an interdigitating suture, and thè angular meets both
thè articular and prearticular. The nature of thè con¬
tact between thè surangular, angular, and articular on
thè mediai surface of thè lower jaw in Tanystropheus
remains elusive (Fig. 53 B).
In thè left lower jaws of MSNM BES SC 265 and
MSNM BES SC 1018 (Fig. 49 C-E) it is confirmed
that thè prearticular formed thè posteriormost part of
thè ventral margin of thè lower jaw without, however,
completely underlying thè retroarticular process. The
prearticular is dorsally delimited from thè articular by
a straight or dorsally slightly concave suture (Figs. 49,
51-52). In MSNM BES SC 1018 there is evidence for
thè prearticular being exposed to a lesser extent than
supposed by Wild on thè mediai surface of thè lower jaw
(Fig. 53).
The coronoid is an element difficult to identify in
thè material of Tanystropheus. Wild (1974: fig. 15,
pi. 18; Fig. 48 in this paper) interpreted as thè right
coronoid in lateral view an isolated element of sub-
triangular shape in PIMUZ T 2484. This element has
a distinct peduncle and an expanded portion entirely
occupied by a concave articular facet. Wild (1974:
37, figs. 15-16; Fig. 53 A in this paper) interpreted
thè peduncle as directed posteriorly and ventrally and
thè coronoid probably overlapping thè surangular,
thè dentary and thè splenial. He also stated that “The
slightly convex dorsal lamina of thè coronoid rises
only insignificantly above thè dentary” (Wild, 1974:
37). This interpretation is difficult to understand. In
my opinion there is no evidence of thè peduncle being
directed ventrally and posteriorly. I think that thè only
way to interpret thè presumed coronoid of PIMUZ T
2484 is to suppose that thè peduncle was directed dor¬
sally and posteriorly, forming a rudimentary coronoid
process, and that thè concave facet on thè expanded
part of thè bone was in contact with thè mediai surface
of thè lower jaw. Based on this interpretation, thè pre¬
sumed coronoid in PIMUZ T 2484 would be thè left
one in lateral view.
In thè new specimens there is no clear evidence
of thè coronoid morphology. However, a rod-shaped
element can be seen dose to thè dorsal margin of thè
mandible in thè left lower jaw of MSNM BES SC 265
(Fig. 49 C). In thè left lower jaw of MSNM BES SC
1018, an approximately rod-shaped but curved ele¬
ment that corresponds in position to thè rod-shaped
element of MSNM BES SC 265 is preserved (Fig.
49 E). A curved isolated element similar in shape and
position to that observed in MSNM BES SC 1018 is
also seen in T. meridensis (Fig. 49 G). This element
was interpreted by Wild (1980a) as an ectopterygoid.
Finally, on thè right lower jaw of MSNM BES SC
1018 (Fig. 53 B), preserved in mediai view, there is a
distinct rod-shaped element similar to that observed in
thè left lower jaw of MSNM BES SC 265 and in thè
same position as all thè others. In thè right lower jaw
of MSNM BES SC 1018 1 tentatively identifìed as part
of thè coronoid also an area ventral to thè rod-shaped
element described above, which might correspond to
thè expanded part of thè isolated bone interpreted by
Wild as a coronoid in PIMUZ T 2484. The rod-shaped
element observed in different specimens in lateral
view might represent thè peduncle of thè same bone,
forming a rudimentary coronoid process.
Dentition
As ascertained by Wild (1974), thè dentigerous bones
in thè small-sized specimens of Tanystropheus are thè
premaxilla, thè maxilla, thè dentary, thè pterygoid, thè
palatine and thè vomer. This is fully confirmed by thè
new specimens, which also confimi thè presence of ante-
rior conical teeth and posterior tricuspid teeth in thè upper
and lower jaws.
Wild considered small-sized specimens to have five
conical premaxillary teeth and thè large-sized ones to
have six (Wild, 1974: 47, fig. 80). However, on thè basis
of thè new specimens there is no difference in thè number
of premaxillary teeth between small- and large-sized
specimens of Tanystropheus. As thè interpretation of thè
premaxillary dentition in MSNM BES SC 1018 is une-
quivocal, thè premaxillary dentition was reconstructed on
thè basis of this specimen.
The left maxilla of MSNM BES SC 1018 bears 14
teeth, all ankylosed. Specimen MSNM BES SC 265
only has 12 tooth positions in thè maxilla. The number
of maxillary teeth documented by Wild in thè small-
sized specimens in thè PIMUZ Collections was 11-13
(Wild, 1974: 47, fig. 80). As Wild counted 12 or pos-
sibly 14 maxillary teeth in thè large-sized specimens,
there is again no appreciable difference in thè number
of maxillary teeth in specimens of different size. While
thè 14 maxillary teeth in MSNM BES SC 1018 are all
tricuspid, thè anteriormost tricuspid maxillary tooth
identifìed in MSNM BES SC 265 is thè fourth (right
maxilla). Wild considered thè anteriormost maxillary
teeth to be conical in thè small-sized PIMUZ specimens
(Wild, 1974: 47, fig. 80). It therefore seems likely that
there is some variability but I based thè reconstruction
of thè maxillary dentition on MSNM BES SC 1018, i.e.
with 14 tricuspid teeth.
The left dentary of MSNM BES SC 1018 bears
19 teeth, with thè anteriormost six conical, and thè
remaining ones tricuspid. In MSNM BES SC 265 thè
anteriormost preserved teeth are conical but thè actual
number of conical and tricuspid teeth, as well as thè
total number of dentary teeth, cannot be ascertained.
Wild (1974: 47, fig. 80) stated that thè small-sized
specimens in thè PIMUZ Collections have at least
eight, or even 11, anterior conical teeth, followed by
tricuspid teeth, for a total of 1 7- 1 9 dentary teeth. Again,
STEFANIA NOSOTTI
62
there seems to be some variability but I reconstructed
thè dentary dentition based on MSNM BES SC 1018.
If, as stated by Wild (1974: 47, fig. 80), thè large-sized
specimens in thè P1MUZ Collections had 19-20? teeth
on thè dentary, there is no appreciable difference in thè
number of dentary teeth in specimens of different size.
The conical teeth of thè upper and thè lower jaw are
interlocking (Ford, 2002), while thè tricuspid teeth of
thè upper jaw are positioned labial to thè corresponding
teeth of thè lower jaw.
At least 12 tooth-attachment sites are present on thè
left pterygoid of MSNM BES SC 1018. Wild (1974:
13-14) counted 14 alveoli in thè pterygoid of PIMUZ
T 2484 (Fig. 43), but emphasized that this number is
variable (he counted less than 13 alveoli in PIMUZ
T 2482). No information can be given regarding thè
shape and size of pterygoid teeth based on thè new
specimens. According to Wild (1974: 13,48) thè ptery¬
goid teeth were all conical and forming a shagreen-like
dentition.
The elements identified as thè left and thè right
palatine in MSNM BES SC 1018 each bear four large
alveoli. Wild (1974: 48) counted five large alveoli
plus a smaller one in thè small-sized specimens in thè
PIMUZ Collections. Eie reported no palatine teeth pre-
served in situ in thè PIMUZ specimens but mentioned
an isolated conical tooth in PIMUZ T 2484 with a
basai diameter matching thè size of thè palatine alveoli
(Wild, 1974: fig. 24 e). This tooth is similar in shape to
thè palatine tooth tentatively identified in MSNM BES
265 (p. 11).
In both MSNM BES SC 1018 and MSNM BES
SC 265 only isolated vomerine teeth were identified.
According to Wild (1974: 47-48) thè total number of
vomerine teeth in thè small-sized PIMUZ specimens
is 9-12. The tiny vomerine teeth identified in thè new
small-sized specimens are similar in shape to those
described by Wild (1974: fig. 24 d). The same is true
for thè vomerine teeth of thè large-sized MSNM V 3663
(p. 38, Fig. 31), some of which are preserved stili in
situ. Based on this specimen thè vomer bore at least ten
alveoli.
The axial skeleton: an overall description
(Figs. 1 , 4-8, 1 4-20, 30, 33-34, 54-60, Pls. I-IV, Tabs. 2-3)
As emphasized by Wild (1974: 52), thè remarkable
compression and/or incomplete preservation of thè ver-
tebral column in thè T. longobardicus material from thè
Grenzbitumenzone (=Besano Formation) hampers thè
detailed reconstruction of vertebral morphology. Speci¬
mens MSNM BES SC 265 and MSNM BES SC 1018
support Wild’s statement and some cruciai issues, such
as thè precise orientation of thè zygapophyseal articular
facets, remain elusive. Thus, as pointed out by Wild
(1974: 52), examination of T. longobardicus specimens is
of little value for understanding thè functional morphol¬
ogy of thè vertebral column, particularly of thè cervical
series. On thè other hand. Wild (1974: 67-94, figs. 39-62)
did describe in detail some isolated, three-dimensionally
preserved vertebrae of T. conspicuus from thè Upper
Muschelkalk of Bayreuth. It was on this material that
he based his arguments for thè mobility of thè vertebral
column and thè posture of thè neck. Maintaining that thè
vertebrae of T. conspicuus were very similar to those of
T. longobardicus. Wild (1974: 52) argued that thè same
conclusions could be applied to both species and recon¬
structed thè vertebrae of Tanystropheus based on T. con¬
spicuus (Wild, 1974: fig. 38).
As discussed below, thè vertebral morphology in
thè new specimens of T. longobardicus is similar to
but not exactly thè same as T. conspicuus. Moreover,
differences in vertebral morphology can be observed
between smaller- and larger-sized individuate of T.
longobardicus in thè PIMUZ Collections. Interestingly,
Dalla Vecchia (2006) recently described isolated, three-
dimensionally preserved vertebrae of Tanystropheus
cf. longobardicus from thè Middle Triassic of northern
Friuli (Italy). These vertebrae belong to large-sized
individuate and some of their features are different
from those described for thè new specimens. These dif¬
ferences in vertebral morphology might alternatively
represent ontogenetic variation within thè same taxon
or taxonomic variation in separate species, and can be
evaluated only with a complete revision of thè genus
Tanystropheus. On one hand, T. longobardicus and T.
conspicuus might be synonymous (Wild, 1 980b: 204;
1987: 39), while thè small- and large-sized T. longo¬
bardicus specimens in thè PIMUZ Collections might
represent two separate taxa (Fraser et al., 2004).
Cervical vertebrae and ribs
(Figs. 1, 7, 14, 30, 34, 54-59, Pls. I-IV, Tab. 2)
All T. longobardicus specimens, those described here
included, confirm that thè cervical vertebral column com-
prises 12 vertebrae (Wild, 1974: 50, although 13 cervicals
were represented by Wild in pi. 1).
The cervical vertebrae three through ten are typi-
cally extremely elongate and slender, and thè increase
in centrum length is not concomitant with an increase in
height.
The length/height ratio can be estimated taking thè
measurement for height at thè middle of thè centrum,
where thè neural spine is absent or weakly developed
as a low keel. In thè new specimens, thè centra of cervi¬
cal vertebrae three through ten are at least nine times as
long as thè height of thè vertebra at thè middle of thè
centrum. The highest ratios occur in cervicals eight and
nine, with centra 13-15 times as long as thè vertebral
height.
In Fig. 54 thè pattern of vertebral lengthening along
thè cervical column in MSNM BES SC 265, MSNM BES
SC 1018 and in thè PIMUZ specimens is compared by
plotting thè ratios “length of cervical vertebrae two (axis)
through twelve / length of thè axis”. The pattern obtained
is similar for thè new specimens and those in thè PIMUZ
Collections, irrespective of thè overall size ot thè indi¬
viduai specimens. As stated by Wild (1974: fig. 29), there
is a peak of vertebral lengthening between thè second and
third cervical vertebrae. After a further, yet moderate,
increase in length between thè third and fourth vertebrae,
vertebral lengthening is negligible between thè tourth and
sixth vertebrae. A graduai, yet continuous, increase in
length is observed again between thè sixth and ninth ver¬
tebrae, thè latter being thè longest (with thè single excep-
TANYSTROPHEVS LONGOBARDICUS. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
63
PIMUZ T 2789
— PIMUZT2779
PIMUZ T 2780
PIMUZ T 2791
— PIMUZ T 2484
♦— PIMUZ T 2795
-t— MSNMBESSC 265
PIMUZ T 2485
•—MSNM BES SC 1018
PIMUZ T 2482
PIMUZ T 2792
-àr- PIMUZ T 2787
— PIMUZT2817
PIMUZ T 2790
PIMUZ T 2819
PIMUZ T 2818
— PIMUZ T 2793
Fig. 54 - Ratios “length of cervical vertebrae two (axis) through twelve/length of thè axis” in Tanystropheus longobardicus (measure-
ments for thè PIMUZ specimens after Wild, 1974: tab. 3). The specimens are listed from thè smallest-sized one to thè largest-sized one.
7
a
idi-
eri
d
ite.
%
tion of PIMUZ T 2818). Posterior to thè ninth vertebra,
thè trend is reversed, with thè vertebral length slightly
decreasing in thè tenth vertebra relative to thè ninth, and
very steeply so between thè tenth and thè 1 2th.
Wild (1974: 52) stated that with increasing age
(=overall length) of thè individuals thè difference in
length between thè eighth, ninth and tenth cervical ver¬
tebrae becomes smaller and smaller but this is apparently
only true for PIMUZ T 2818, in which cervical vertebrae
eight through ten reach very similar lengths, thè eighth
being thè longest. Wild (1974: 52) also maintained that
in specimens over 2 meters overall length (i.e. his “adult”
individuals) thè mutuai rate of lengthening of thè cervical
vertebrae changes. In particular, with increasing overall
length, thè difference in length of vertebrae four through
ten would decrease, because of thè positive allometric
growth in thè third and fourth vertebrae, and slower or
no additional growth in thè ninth vertebra. However,
Wild’s statement is not confirmed by thè pattern shown
in Fig. 54, which shows instead that thè mutuai rate of
lengthening of thè cervical vertebrae is not affected by thè
overall length of thè specimens. I think that thè pattern
of lengthening of thè cervical vertebrae in specimens of
different overall length does not contradict thè hypothesis
that small- and large-sized specimens of Tanystropheus
represent different ontogenetic stages of thè same species.
However, contra Wild (1974: 52), I do not think that it
supports thè hypothesis either.
Both in thè PIMUZ material and in thè new specimens
thè atlas is never preserved as a complete element. The
atlas-axis complex of T. longobardicus was reconstructed
by Wild (1974: fig. 31) on thè basis of thè large-sized
specimen PIMUZ T 2819 (Fig. 55). This specimen pre-
serves thè paired neural arches of thè atlas, and other
isolated elements interpreted by Wild as thè proatlas, thè
atlas centrum, and thè atlas intercentrum (Wild, 1974:
fig. 6, pi. 16). The right atlas neural arch of PIMUZ T
2819 is very similar in shape to thè left atlas neural arch
of MSNM BES SC 1018 (Figs. 10-12, 56 B). Both are
exposed in lateral view. The neural arch is an L-shaped
element with a posterior and a ventral ramus. In PIMUZ
T 2819 thè posterior ramus is more developed than thè
ventral one, while in MSNM BES SC 1018 thè two rami
approximately reach thè same length. The postzygapoph-
ysis takes thè form of a rib-shaped, horizontal relief pro-
jecting backwards from thè posterior ramus of thè neural
arch. According to Wild (1974: 55, fig. 31), thè neural
arch projected anteriorly into a pointed process, thè cornu
at/anticum but thè presence of this process on thè left atlas
neural arch of PIMUZ T 2819 is not clear. By contrast,
I assume that thè right atlas neural arch of MSNM BES
SC 1018 is articulated anteriorly with thè proatlas (Figs.
10-12). Based on this assumption it is unlikely that thè
two isolated elements interpreted by Wild as thè proatlas
in PIMUZ T 2819 (Fig. 55) were correctly identified. As
an alternative I suggest that thè element labelled “pa?”
in Fig. 55 and partially overlapped by thè left quadrate is
thè proatlas in articulation with thè right atlas neural arch.
Wild considered this fragment of bone to be part of thè
left quadrate. I do concur with Wild on thè identification
of thè atlas centrum and intercentrum in PIMUZ T 2819
(see also Figs. 10-12).
64
STEFANIA NOSOTT1
Fig. 55 - Tanystropheus longobardicus, PIMUZ T 2819, atlas and axis on thè interpretation of Wild, 1974. pa?: proatlas on thè inter-
pretation of thè author. Scale bar 20 mm. Photo: Heinz Lanz, PIMUZ.
The morphology of thè axis is well documented in
thè known material of Tanystropheus, and is apparently
thè same in both small- and large-sized specimens (Wild,
1974: figs. 30-31). The reconstruction given by Wild
(1974: fìg. 31), based on thè large-sized PIMUZ T 2819
(Fig. 55), is consistent with thè new specimens (Fig. 56).
Apart from thè atlas, thè axis is invariably thè shortest
vertebra of thè cervical column. The articular surfaces
of thè centrum are not inclined. The anterior surface is
ventrally bevelled, receiving an elongate intercentrum. A
latero-ventral margin (die Lateroventralkante or margo
inferior, sensu Tschanz, 1986: 60) running along thè
entire length of thè centrum delimits a well developed
ventral keel (=hypapophysis sensu Wild, 1974: 55). As
in thè subsequent cervicals (see below), thè latero-ventral
margin bifurcates anteriorly into two sharp cristae for thè
rib articulation. As compared to that of thè subsequent
cervicals, thè neural arch of thè axis is well developed,
raising anteriorly into a rounded neural spine. It has an
anterior tubercle for thè insertion of thè atlanto-occipital
ligaments (Wild, 1974: 56; Fig. 56 A in this paper). The
prezygapophyses are apparently short and stocky (Wild,
1974: fig. 31) and do not project bevond thè anterior end
of thè centrum. According to Wild (1974: 56), they origi-
nally had horizontal articular surfaces but this cannot be
verifìed in thè new specimens. The articular relationship
between thè atlas neural arch and thè axis, as interpreted
by Wild (1974: fig. 31), is confirmed by MSNM BES SC
265 (p. 13, Fig. 56 A). An articular facet at thè basis of thè
axis neural arch, as seen in PIMUZ T 2819 (Fig. 55) prob-
ably received thè atlas neural arch. The postzygapophyses
of thè axis are elongate and project beyond thè posterior
end of thè centrum.
The morphology of cervicals three through ten is simi-
lar in thè new specimens, and it is particularly clear in thè
more three-dimensionally preserved vertebrae of MSNM
BES SC 1018. The anterior and posterior articular surfaces
of thè centra are not inclined. By contrast, Wild (1974: 95) :
ascertained that in T. conspicuus these surfaces are inclined
anteriorly to a variable degree. As is typical for Tanystroph¬
eus , strongly keeled margins run along thè centrum (Wild,
1974; Tschanz, 1985, 1986). Flowever, a latero-dorsal
margin (die Laterodorsalkante or margo lateralis, sensu
Tschanz, 1986: 61) running along thè entire length of thè
centrum and continuing onto thè dorsal margins of thè pre-
and thè post-zygapophysis was not observed in thè new
specimens. The neural spine is poorly developed.
The third and fourth cervicals of MSNM BES SC 1018
(Fig. 57 A) have a strongly keeled latero-ventral margin
running along thè entire length of thè centrum, approxi-
mately at its half-height. On this basis thè cross section in
thè middle of thè centrum would have been sub-circular.
The latero-ventral margin is posteriorly bifid. Anteriorly it
bifurcates into two sharp cristae separated by a groove, tor
thè rib articulation. This latter feature is very distinct also
in thè cervical vertebrae of MSNM BES SC 265 (Fig. 58).
TANYSTROPHEUS LONGOBARDICUS. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
65
Fig. 56 - Tanystropheus longobardicus , axis. A) MSNM BES SC 265,
right lateral view. Scale bar 5 mm. B) MSNM BES SC 1018, lefìt lateral
view. Scale bar 5 mm. Photos: Massimo Demma.
The dorsal crista continues onto thè ventral margin of thè
prezygapophysis. A keeled ventral margin {die Ventral-
kante or margo ventralis, sensu Tschanz, 1986: 59) was
probably not developed in these vertebrae. The neural
spine is only anteriorly developed in thè third cervical,
where it consists of a low ridge projecting into a pointed
tip rising up from thè dorsal surface of thè centrum. In thè
fourth cervical it is rudimentarily developed also posteri-
orly. Anteriorly, thè neural spine bears a sharply pointed
process, and ventral to it is a sub-triangular fossa bordered
by raised margins.
In MSNM BES SC 1018, posterior to thè fourth
cervical, thè raised latero-ventral margin runs along thè
centrum from thè prezygapophysis, becoming lateral to
thè fiat ventral surface of thè centrum, and levels out in
its posterior part (Tschanz, 1986: 60) (Fig. 57 B). On this
basis thè cross section in thè middle of thè centrum would
have been sub-triangular (Tschanz, 1986: 60). A ventral
keel seems to be developed anteriorly in cervicals eight
through ten. In cervicals five through ten thè neural spine
extends along thè entire length of thè centrum as a con-
tinuous, yet very low, dorsal keel, which becomes higher
anteriorly and posteriorly (Fig. 57 B).
In thè specimens of T. longobardicus of thè PIMUZ
Collections Wild (1974: 58) observed that from thè sixth-
seventh cervical on backwards thè neural spine grows sig-
nificantly higher anteriorly and is gradually reduced pos¬
teriorly. In cervicals three through ten of thè larger speci¬
mens a dorsal keel is absent, and thè neural spine is almost
completely reduced posteriorly (Tschanz, 1986: 62). A
neural spine only anteriorly developed was described
by Dalla Vecchia for thè large, three-dimensionally pre-
served cervical MFSN 31579 {T. cf. longobardicus. Dalla
Vecchia, 2006: fig. 8), tentatively identified as thè ninth
(Dalla Vecchia, 2006: fig. 17). Dalla Vecchia suggested
that this feature might be an apomorphy indicating that
thè new material from Friuli represents a new species of
Tanystropheus.
ns pzp
Fig. 57 - Reconstruction of an anterior cervical vertebra (A) and of a mid-cervical vertebra (B) in small-sized specimens of Tanystro¬
pheus longobardicus. Left lateral view. Not to scale. Watercolor: Massimo Demma.
66
STEFANIA NOSOTTI
Fig. 58 — Tanystropheus longobardicus , MSNM BES SC 265, articula-
tion between thè fifth and sixth cervical vertebrae. Scale bar 5 mm.
Photo: Massimo Demma.
Both pre- and postzygapophyses of thè cervical verte¬
brae three through ten project well beyond thè anterior and
posterior ends of thè centrum. A strongly keeled margin
delimits thè postzygapophysis ventro-laterally, extending
anteriorly up to approximately one half thè length of thè
centrum. In MSNM BES SC 1018 thè postzygapophyses
have a finely striated surface for muscular insertion {die
Intertransversal-Muskulatur, Wild, 1974: 58).
In MSNM BES SC 265 and MSNM BES SC 1018,
thè 1 lth vertebra (Fig. 14) is similar in its generai shape to
that fìgured by Wild (1974) in fig. 97 a (PIMUZ T 1270).
The centrum is remarkably shorter than that of thè preced-
ing cervicals, and thè neural arch higher, yet not as high
as in thè corresponding cervical of T. conspicuus (Wild,
1974: fig. 97 b-c). The anterior and posterior articular
surfaces of thè centrum are not inclined. Two distinct
facets for thè rib articulation are bome on two tubercles
positioned antero-ventrally on thè centrum and separated
by a groove. A latero-ventral margin runs from thè dorsal
tubercle along approximately three-fourths ofthe centrum,
delimiting a ventral keel. Both pre- and postzygapophyses
project beyond thè articular surfaces of thè centrum. The
neural spine runs along thè entire length ot thè centrum as
a low keel, raising anteriorly and posteriorly, so that thè
outline of thè neural spine in lateral view is concave. As
is also thè case in thè preceding cervicals, thè neural spine
deepens anteriorly into a sub-triangular fossa bordered by
raised margins.
In MSNM BES SC 265 (Figs. 4, 7) thè 12,h cervi¬
cal is very short, thè length of its centrum only slightly
exceeding that of thè axis (Tab. 2). Albeit damaged, its
neural arch is clearly much higher than in thè preceding
cervicals. As emphasized by Renesto (2005), thè poste¬
rior articular surface of thè centrum is antero-posteriorly
inclined. Similar to thè llth vertebra, two antero-ventral
tubercles separated by a groove bear distinct facets for thè
rib. A latero-ventral margin runs from thè dorsal tubercle
along almost thè entire length of thè centrum, delimiting
a ventral keel. By contrast with thè postzygapophyses, thè
prezygapophyses project only very slightly beyond thè
anterior end of thè centrum. This features are consistent
with thè description given by Wild (1974: 80, fig. 50) for
thè 12th cervical of T. conspicuus.
Little can be added to our understanding of thè articu¬
lation of thè cervical vertebrae on thè basis of thè new
specimens. As expected, thè prezygapophyses are dor-
sally overlapped by thè postzygapophyses. Observation of
thè disarticulated vertebrae four and five in MSNM BES
SC 1018 (Fig. 59) reveals that thè postzygapophyseal
articular facet was probably sub-vertical in this region.
The postzygapophyseal articular facet of thè fourth cervi¬
cal has an ovai, antero-posteriorly elongate shape and is
positioned ventral and mediai to thè postzygapophyseal
process ( sensu Rieppel, 2001). The latter projects pos¬
teriorly beyond thè articular facet, roofing it, and in thè
articulated vertebrae would lie on a sub-horizontal shelt
bome on thè prezygapophysis. This shelf is seen on thè
fifth cervical, mediai to thè dorso-lateral margin of thè
prezygapophysis.
In thè large, three-dimensionally preserved cervical
MFSN 31579 (Dalla Vecchia, 2006: 37, fig. 8) thè postzi-
gapophyseal facets slant in a dorso-lateral/ventro-medial
piane, forming an angle of 45° with thè mediai piane of
symmetry. The prezygapophyseal facets appear to be
vertical in MFSN 31579 but they face dorso-medially in
other cervicals comprised in thè material studied by Dalla
Vecchia (2006: 37-38).
In both MSNM BES SC 265 and MSNM BES SC
1018, thè cervical ribs dump in tight, paired bundles
ventral to thè cervical column. Both specimens do not
add new information on thè well-known morphology of
thè cervical ribs in Tanystropheus. The head of each rib
in dorsal view is axe-shaped, with an anterior and poste¬
rior process. The latter is present also in thè anteriormost
cervical ribs.
Wild (1974: 98) emphasized that thè articular facets
for thè ribs bome on thè centmm are different between
T. conspicuus and T. longobardicus. He described thè
Fig. 59 — Tanystropheus longobardicus , MSNM BES SC 1018, articu¬
lation between thè fourth and fifth cervical vertebrae. Scale bar 5 mm.
Photo: Massimo Demma.
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
67
articular facets of thè latter as elliptical and positioned
in grooves. According to Wild (1974: 59) thè tuberculum
and capitulum in thè first eight ribs posteriorly formed
a single synapophysial articular surface, these ribs thus
being functionally olocephalous. This cannot be con-
fìrmed in thè new specimens. However, an isolated rib
head preserved dose to thè tenth cervical vertebra in
MSNM BES SC 1018 exhibits a completely separated
tuberculum and capitulum. The llth cervical of thè same
specimen unequivocally has two articular facets for thè
rib (Fig. 14), as do thè llth and 12th cervicals in MSNM
BES SC 265 (Figs. 4, 7). In thè latter specimen, thè tuber¬
culum and capitulum of thè 1 2th cervical rib are distinctly
shorter and higher, as is typical of a dichocephalous rib
(Fig. 4).
In thè specimens described here, as mentioned
above, thè tuberculum and capitulum of thè function¬
ally holocephalous ribs articulate on two divergent crests
- separated by a deepened triangular surface - positioned
ventro-laterally on thè centrum. In MSNM BES SC 265,
this feature is clearly seen in vertebrae of thè series two
through seven (Fig. 58 A). In MSNM BES SC 1018, it
can be clearly seen in thè third and fourth vertebrae. In thè
sixth vertebra thè articulation of thè rib with thè centrum is
apparently more ventral. In fact, moving posteriorly there
seems to be a progressive ventral shift in thè position of
thè rib articulation. The articular area on thè centrum in
thè mid- and posterior cervicals cannot be described on
thè basis of MSNM BES SC 1018.
The extremely thin and elongate shafts of thè cervical
ribs are closely packed in MSNM BES SC 265, and it is
difficult to ascertain either their length or number in thè
different regions of thè cervical vertebral column. The
same is true for MSNM BES SC 1018, in which thè shafts
of thè cervical ribs are also extensively broken. In MSNM
BES SC 265 one of thè fourth ribs is isolated, and its shaft
apparently intact. Its length exceeds that of three cervical
vertebrae.
Dorsal vertebrae and ribs
(Figs. 1, 4-5, 7, 15-18, 20, 33, 60, Pls. I-IV)
All thè known specimens confimi that thè dorsal verte¬
bral column of T. longobardicus consists of 1 3 vertebrae.
However, Wild (1974) distinguished 11 dorsal vertebrae
and two “lumbar” vertebrae. Indeed, thè morphology of
thè last two dorsals is different from that of thè preceding
ones (see bolow).
Wild (1974) described thè dorsal vertebrae in T. longo¬
bardicus on thè basis of thè large-sized specimen PIMUZ
T 2817 and thè smaller PIMUZ T 1270. He maintained
that in thè other specimens in thè PIMUZ Collections
these vertebrae were too badly crushed to permit a recon-
struction.
Common features of thè dorsal vertebrae are thè ven-
trally concave centra and tali neural spines, as compared
to those of thè cervical vertebrae. The neural spines are
dorsal ly flattened and reinforced by a stout, rough ridge
for muscular insertion.
The dorsal vertebrae can be grouped into an ante-
riormost region, comprising vertebrae one through
three; an intermediate region, comprising vertebrae
five through ten; and a posterior or “lumbar” region
comprising thè last two dorsals. The fourth and llth
dorsal vertebrae cannot be clearly assigned to anyone
of these regions (see below).
In Fig. 60 A thè anteriormost dorsal vertebrae are
reconstructed on thè basis of MSNM BES SC 265 (Fig. 4)
and MSNM BES SC 1018 (Fig. 16). The height meas-
ured from thè ventral surface of thè centrum to thè top of
thè neural spine is approximately equal to thè length of
thè centrum. The centrum is deeply concave and keeled
ventrally. The posterior articular surface slants antero-
posteriorly. Typically, thè anteriormost dorsal vertebrae
bear two articular facets for dichocephalous ribs. The
wide, sub-circular articular facet for thè capitulum of
thè rib is bome on a distinct tubercle positioned ventrally
and anteriorly on thè centrum. A rounded articular facet
for thè tuberculum is bome on a stout but short trans¬
verse process. The transverse process is positioned on
thè anterior half of thè centmm, where thè neural arch
is fused to thè centmm but is somewhat more ventral
Fig. 60 - Reconstruction of an anteriormost dorsal vertebra (A)
and of a mid-dorsal (B) in small-sized specimens of Tanystropheus
longobardicus. Right lateral view. Not to scale. Watercolor: Massimo
Demma.
68
STEFANIA NOSOTTI
in thè first dorsal. A stout bony bridge extends between
thè transverse process and thè prezygapophysis. A short
buttress supports thè transverse process ventrally, both
anteriorly and posteriorly. This arrangement is similar to
that described by Wild (1974: 61) for thè first five dorsals
in T. ìongobardicus. The prezygapophyses are short and
project dorsally and only slightly beyond thè anterior end
of thè centrum. The postzygapophyses are located high
on thè neural arch, separated from thè centrum by a deep
notch. They do not project beyond thè posterior end of thè
centrum. The orientation of thè zygapophyseal articular
facets is sub-vertical. The neural spine takes thè form of
a truncated pyramid, with anteriorly and posteriorly pro-
jecting processes at thè top and a fiat dorsal surface. It is
approximately as long as tali.
The morphology of thè anteriormost dorsal verte-
brae in MSNM BES SC 265, MSNM BES SC 1018,
and thè small-sized T. ìongobardicus specimens in thè
PIMUZ Collections (Peyer, 1931: pls. 1-3) differs from
that reconstructed by Wild (1974: fìgs. 38 c, 52) for T.
conspicuus. In thè latter species, thè height of thè anteri¬
ormost dorsals, measured from thè ventral surface of thè
centrum to thè top of thè neural spine, is approximately
twice thè length of thè centrum. The neural spines are
sub-rectangular in shape and taller than long (Renesto,
2005: 378).
The centrum of thè mid-dorsal vertebrae (Fig. 60 B) is
poorly preserved both in MSNM BES SC 265 (Fig. 7) and
MSNM BES SC 1018 (Figs. 15, 20) but on thè basis of thè
material in thè PIMUZ Collections it is ventrally concave
(see also Renesto, 2005: fig. 4; Dalla Vecchia, 2006: fig.
10). The length of thè centrum is equal to or exceeds thè
height measured from thè ventral surface of thè centrum
to thè top of thè neural spine. Neither thè anterior nor
posterior articular surfaces of thè centrum are inclined.
The single, sub-ovai articular facet for an holocephalous
rib is bome on a short transverse process. However, it
might look shorter than it really was since thè specimens
have been greatly crushed (see for comparison specimen
MCSN 4451, Renesto, 2005: fig. 4 A). The transverse
process is positioned dose to thè base of thè neural arch,
approximately mid way along thè centrum. A short, ridged
buttress supports thè transverse process antero-ventrally
(centrodiapophyseal lamina sensu Dalla Vecchia, 2006).
Short prezygapophyses, positioned at thè same level as thè
transverse processes, project very slightly beyond thè end
of thè centrum. The lateral margin of each prezygapophysis
takes thè form of a laterally projecting crest reaching thè
transverse process posteriorly (Figs. 15, 20, 60 B). The
postzygapophyses are located low on thè neural arch and
project only slightly beyond thè end of thè centrum. The
orientation of thè zygapophyseal articular facets is inter-
preted as sub-horizontal (see also Tschanz, 1985: 174). The
neural spines are sub-rectangular and approximately twice
as long as tali (Renesto, 2005: 378).
The morphology of thè mid-dorsals of thè new speci¬
mens from Besano is thus different in some respects from
that of thè mid-dorsals of T. conspicuus (Wild, 1974: figs.
38 d-e, 54) and of other large-sized specimens of Tanys-
tropheus such as PIMUZ T 2818 {T. ìongobardicus. Wild,
1974: pi. 13) and MFSN 31596 ( T . cf. ìongobardicus.
Dalla Vecchia, 2006: fig. 10). In these large dorsals thè
height measured from thè ventral surface of thè centrum
to thè top of thè neural spine is twice or more thè length of
thè centrum. The neural spine is antero-posteriorly short
and approximately as long as tali. This was linked by
Wild (1974: 61) to thè need for larger and higher insertion
areas for thè musculature in thè large-sized individuate.
A diapoprezygapophyseal lamina {sensu Dalla Vecchia,
2006) runs obliquely from thè transverse process on thè
lateral side of thè prezygapophysis. The transverse proc¬
ess is rather high on thè neural arch. Specimen MFSN
3 1 596 has quite long and dorso-ventrally flattened trans¬
verse processes with antero-posteriorly expanded ends,
which according to Dalla Vecchia (2006: 43) might be an
apomorphy of thè taxon represented by thè material from
Friuli.
Specimen MSNM BES 215 described here (pp. 38-
39) ateo represents a large-sized individuai. The short
postzygapophyses, positioned low on thè neural arch
and provided with sub-horizontal articular facets, sug-
gest that this vertebra is in all probability a mid-dorsal.
As in MSNM BES SC 265 and MSNM BES SC 1018, thè
transverse process projects laterally from thè area where
thè neural arch meets thè centrum. By contrast, although
thè neural spine of MSNM BES 215 is broken, thè height
of its preserved part suggests that it was very tali. Finally,
this dorsal vertebra is unique for all known Tanystropheus
material for thè presence of postzygapophyseals canate in
thè postzygapophyseal trough. Wild (1974: 81) described
a deep postzygapophyseal trough {die postzygapophyseale
Grube) and postzygapophyseal canate {der Doppelkanal',
das Ròhrenpaar sensu von Meyer, 1847-1855) in thè cer-
vical vertebrae of T. conspicuus. Rieppel (2001) described
a similar feature in thè cervical vertebrae of T. haasi. In
this species, postzygapophyseal grooves in thè floor of thè
postzygapophyseal trough extend far anteriorly into thè
neural arch, separated from one another by a thin vertical
bony septum, and separated from thè neural canal by an
equally thin horizontal septum. As formerly pointed out
by Wild (1974: 81) thè postzygapophyseal canate end
blindly. Wild (1974: 81) stated that a postzygapophyseal
trough could be seen ateo in thè dorsal, lumbar and caudal
vertebrae of Tanystropheus. However, postzygapophyseal
canate have been never described in thè post-cervical
vertebrae.
Based on thè small-sized Tanystropheus specimen
MCSN 4451 Renesto (2005: fig. 4) assumed that thè
nature of thè rib articulation changes gradually along
thè dorsal vertebral series and described an intermedi¬
ate condition in which two articular facets for thè rib are
very dose to each other but stili not confluent. Specimen
MSNM BES SC 265 unequivocally shows that from thè
fifth dorsal on backwards there is a single articular facet
for thè rib, bome on thè transverse process. A dose look
at MCSN 445 1 reveals that thè ventral articular facet on
thè vertebra figured by Renesto (2005) in fig. 4 A is in fact
an artefact of fossilization, and that there is but a single
articular facet for thè rib. I was not able to confirm thè
presence of two articular facets dose to each other in thè
vertebra figured by Renesto (2005) in fig. 4 B. Indeed
thè vertebra is badly crushed and difficult to interpret.
In MSNM BES SC 265 thè fourth dorsal is very poorly
preserved. However, as far as can be judged from thè
morphology of thè transverse process and prezygapo¬
physis, it has a single articular facet for thè rib just like
all subsequent dorsals. By contrast, on thè basis of speci¬
mens PIMUZ T 2817 and PIMUZ T 2787, Wild (1974)
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
69
considered thè fourth dorsal had two articular facets for
thè rib, while from thè fifth on there was a single articular
facet. An intermediate condition similar to that described
by Renesto (2005) for MCSN 445 1 might be present in an
isolated dorsal of MSNM BES SC 1018, that I tentatively
interpret as thè fourth (Fig. 17). The strongly concave and
keeled centrum, thè antero-dorsally projecting prezygap-
ophysis, and thè shape of thè neural spine are typical char-
acters of thè anteriormost dorsal vertebrae. By contrast,
thè postzygapophysis is very similar in shape to that of
thè subsequent dorsals, probably hearing a sub-horizontal
articular facet. This vertebra has an unequivocal single
articular facet for thè rib on thè transverse process, with
a stout, oblique, and distinctly projecting crest extending
between thè transverse process and thè prezygapophysis.
Another crest extends from thè transverse process to thè
antero-ventral area of thè centrum. Along this crest there
may be a second articular facet. If this is thè case, this
vertebra has two facets for thè rib articulation showing a
condition similar to that described by Renesto (2005). I
conclude that there is individuai variation in thè position
of thè facets for thè rib articulation on thè anteriormost
dorsal vertebrae. However, as a rule thè mid-dorsal verte¬
brae typically have a single facet bome on thè transverse
process.
The morphology of thè llth dorsal vertebra is dif-
ferent from both preceding and subsequent vertebrae.
Although this vertebra is not well preserved in MSNM
BES SC 265 (Fig. 5), it is very similar to an isolated,
well preserved vertebra in MSNM BES SC 1018 (Fig.
18). The centrum of thè llth dorsal bears a single, wide,
roundish facet for thè rib articulation bome on a stout
and short transverse process. The transverse process is
positioned very dose to thè prezygapophysis at thè base
of thè neural arch. Both anteriorly and posteriorly, thè
transverse process is supported by short ventral but-
tresses. The anterior buttress is strongly ridged. The
prezygapophysis is shaped as in thè mid-dorsal verte¬
brae, with a presumed sub-horizontal articular facet. The
postzygapophysis is positioned high on thè neural arch,
and in shape is more like thè anteriormost dorsals than
thè mid-dorsals.
Dorsal vertebrae 12 and 13 (“lumbar vertebrae”), are
poorly preserved in MSNM BES SC 265 (Fig. 5) but are
strikingly different from thè preceding ones. The neural
spine is square and distinctly shorter than in thè mid-
dorsals. The postzygapophysis is positioned high on thè
neural arch. Finally, these vertebrae bear long pleurapo-
physes1.
In both MSNM BES SC 265 and MSNM BES SC
1018, thè dorsal ribs are mostly disarticulated and
scattered, and many of them are missing alltogether.
Observation of thè preserved elements suggests that thè
morphology of thè dorsal ribs in thè new specimens is
similar to that described by Wild (1974: 63, fig. 35). In
MSNM BES SC 265, thè first dorsal rib is preserved in
association with thè first dorsal vertebra (Fig. 4). The
tuberculum is short and thick, thè capitulum is longer
and more slender. The shaft is distally overlapped by
thè second dorsal vertebra so that its full length cannot
be ascertained. An anteriormost, isolated dichocephal-
ous rib is also preserved in MSNM BES SC 1018 (p.
28), which is shaped like thè first dorsal rib of MSNM
BES SC 265. Its shaft does not seem broken, although
it appears rather short. The second dorsal rib is not pre¬
served in MSNM BES SC 265 but thè third is. It is not
completely articulated with thè third vertebra but thè
tuberculum and thè capitulum He very dose to thè cor-
responding vertebral articular facets. This rib is slender
with a short tuberculum and a longer capitulum. The
shaft is gently curved. In thè new specimens it is not
possible to ascertain whether thè fourth dorsal rib was
stili dichocephalous but on thè basis of thè morphology
of thè fourth dorsal vertebra in MSNM BES SC 265 (see
above) it probably was not.
Sacrai vertebrae
(Figs. 1,6, 8, 19, Pls. I-IV)
The sacrai vertebrae are readily identifìed based on
their expanded pleurapophyses1 that articulated with thè
ilium. Specimen MSNM BES SC 265 (Figs. 6, 8) con-
firms that there are two sacrai vertebrae in Tanystropheus .
The isolated sacrai vertebra preserved in MSNM BES
SC 1018 (Fig. 19) is interesting, because it display s thè
vertebra in lateral view. In MSNM BES SC 1018, thè
overall shape of thè vertebra is quite different from that
of thè sacrai vertebrae described by Wild for T. longo-
bardicus (Wild, 1974: 62) and T. conspicuus (Wild, 1974:
figs. 55, 56, tab. 6). These display a shorter, less slender
centrum and a tali, sub-rectangular and antero-posteriorly
short neural spine. A large, three-dimensionally preserved
sacrai vertebra referred to Tanystropheus cf. longo-
bardicus (MFSN 31552) was recently described by Dalla
Vecchia (2006: fig. 11). It does not preserve thè neural
spine but thè centrum is very similar to that of thè sacrals
in T. conspicuus.
Caudal vertebrae
(Figs. 1,6, Pls. I-IV, Tab. 3)
The description of thè caudal vertebrae is based on
MSNM BES SC 265, which preserves thè caudal series in
its originai proximo-distal sequence (Figs. 1, 6). The iso¬
lated caudals in MSNM BES SC 1018 do not add relevant
information and neither specimen provides any reai detail
of thè caudals distai to thè 1 lth (Fig. 1, Pls. I-IV).
The centra of thè first three caudal vertebrae are
approximately as long as those of thè second and thè
1 lth dorsal vertebrae. The length of thè subsequent centra
steadily increases proceeding distally along thè caudal
series (Tab. 3). Up to thè eight vertebra, thè centra are
1 Wild (1974: 62) used thè term “pleurapophysis” to indicate a rib fused to thè transverse process. As emphasized by Romer (1956:
276), once a rib is completely fused to thè transverse process without a discemible suture between thè two, it is difficult to establish
whether it represents a rib fused to thè transverse process or an elongate transverse process. Both Renesto S. (pers. comm., 2006)
and I observed that there is sometimes a line of separation visible at thè very base of thè wall of thè neural arch, without even a short
transverse process (see for example thè third caudal in MSNM BES SC 265). However, it is difficult to teli whether it is a suture or a
fracture. in thè absence of additional arguments clarifying this issue, I will adopt thè term “pleurapophysis” as used by Wild.
70
STEFANIA NOSOTTI
strongly concave ventrally. Caudals one through seven
have well developed latero-ventral margins, delimiting
a ventral keel in thè sixth and seventh. The articular sur-
faces of thè centra are not inclined.
The first neural arch that can be observed is that of thè
sixth caudal. The neural spine is lower than in thè mid-
dorsals, and dorsally flattened. Distai to thè sixth vertebra,
thè neural spine progressively shortens and shifts posteri-
orly. The pre- and postzygapophyses lie at approximately
thè same level. The prezygapophyses project well beyond
thè anterior end of thè centrum, while thè postzygapo¬
physes are shorter, extending only to approximately thè
posterior end of thè centrum. A crest connects thè pre-
and postzygapophyses on thè lateral side of thè neural
arch. This crest is very well developed in vertebrae seven
through nine but reduced in thè tenth vertebra and almost
absent in thè 1 lth. The zygapophyseal articular surfaces
appear sub-horizontal (see also Tschanz, 1985: 174; Dalla
Vecchia, 2000: fìg. 2; 2006: fig. 12).
In MSNM BES SC 265, thè first through eighth
caudals display pleurapophyses (see note on p. 69), that
increasing in length from thè first to thè third element in
thè caudal series. The pleurapophyses of thè fourth and
fìfth caudals are broken and cannot be measured. A frag-
ment of one such pleurapophyses (PI. II) suggests that
they were stili rather long. Distai to thè fifth vertebra, thè
length of thè pleurapophyses is significantly reduced. In
thè eighth caudal thè pleurapophysis is rudimentary, and
there are no pleurapophyses on thè subsequent caudals.
The number of caudal vertebrae with more or less well
developed pleurapophyses is therefore consistent with thè
number given by Wild (1974: 62) of seven-eight for thè
small-sized specimens of T. longobardicus . As pointed
out by Wild, thè position of thè pleurapophysis on thè
centrum shifts diagonally ventrally and posteriorly pro-
ceeding distally along thè caudal series. The pleurapophy¬
ses of thè proximal caudal vertebrae are stoutly built (see
also thè proximal caudals in MSNM BES SC 1018, Pls.
III-IV), as shown by their wide antero-posterior base, and
they are strongly dorso-ventrally flattened.
None of thè preserved caudal vertebrae in thè new
specimens is similar in shape to thè proximal caudals of
T. conspicuus (Wild, 1974: figs. 38 f, 57-61). The latter
possess very tali and antero-posteriorly short, sub-rectan-
gular neural spine. The postzygapophysis is very short
and apparently bears an oblique articular facet. However,
given thè rather poor preservation of caudals one through
five in MSNM BES SC 265 and of thè proximal isolated
caudals in MSNM BES SC 1018, thè overall shape of
these vertebrae cannot be described and compared to thè
specimens in thè PIMUZ Collections. The shape of thè
sixth caudal vertebra in MSNM BES SC 265, and ot those
distai to it, is more like that of thè mid-caudal vertebra of
T. conspicuus in Wild’s fig. 62. Dalla Vecchia (2000: fìg.
2, MFSN 25761; 2006: fig. 12, MFSN 31549) described
proximal isolated caudal vertebrae of Tanystropheus ffom
thè Middle Triassic of northem Italy, belonging to rather
large individuate. The shape of thè neural spine of these
vertebrae is ateo similar to that described by Wild for thè
proximal caudate of T. conspicuus.
The first preserved Chevron in MSNM BES SC 265 is
that between thè seventh and eighth vertebrae, and thè last
between thè 11* and thè 12*. As stated by Wild (1974: 67,
fig. 37), thè shape of thè chevrons changes moving dis¬
tally down thè vertebral series. Their ventral rami become
shorter, and antero-posteriorly expanded in lateral view. An
isolated Chevron is preserved in MSNM BES SC 1018 (p.
29, Pls. III-IV), showing that thè two branches of thè chev¬
rons meet ventrally, forming a median spine, and are dor¬
sally connected by a transverse bar in a condition slightly
different from that shown in Wild’s (1974) fig. 67.
In MSNM BES SC 265, thè vertebral series distai to i
thè 1 lth caudal vertebra is complete but it cannot be ascer-
tained how many vertebrae are present. On thè basis of its
length (p. 1 5), thè series apparently comprises a minimum
of 30 vertebrae of a length comparable with that of thè
proximal caudate but this number was surely far higher
if thè progressive shortening of thè centra towards thè
tip of thè tail is taken into account. In fact, thè estimated
length of thè distalmost caudal vertebrae of MSNM BES
SC 265 is 2.5 mm. Unfortunately, thè caudal series is not
complete and poorly preserved in MSNM BES SC 1018.
According to Wild (1974: 62), thè overall number of
caudal vertebrae was over 50.
The presence of fracture planes indicative of caudal
autotomy (Wild, 1974: 94) cannot be confirmed on thè
caudal vertebrae of MSNM BES SC 265.
The mobility of thè vertebral column in Tanystropheus
The description of thè pre- and postzygapophyseal
articular facete in thè three-dimensionally preserved ver¬
tebrae of T. conspicuus (Wild, 1974: 96-98) remains thè
best starting point for any understanding of thè mobility
of thè vertebral column in Tanystropheus.
At present, thè only detailed descriptions and interpre-
tations of thè axial locomotory System in Tanystropheus
are those of Wild (1974) and Tschanz (1985; 1986; 1988).
These authors, however, carne to different conclusions.
Analysing thè shape and thè orientation of thè zygapo¬
physeal facets, and evaluating thè role played by thè very
thin and elongate cervical ribs in thè mechanics of thè cer-
vical vertebral column, Wild (1974) asserted that thè neck
of Tanystropheus was very mobile and, when on land,
could assume an S-shaped configuration.Wild recognized
three distinct regions in thè neck. In thè anterior region,
he considered sagittal movements predominated but with
some lateral flexion. In thè mid-region, sagittal flexion
predominated, while in thè caudal region both sagittal and
lateral flexion were possible.
Wild (1974) ateo stated that, compared to thè cervical
series, thè mobility of thè dorsal and caudal series was
less. He assumed that thè anteriormost dorsal column
had thè capability to flex dorsally and laterally but pro-
ceeding posteriorly towards thè sacrai region thè overall
mobility of thè dorsal vertebral column decreased. In thè
caudal region as a whole he considered dorsiflexion to be
reduced and to be absent alltogether in thè proximal cau¬
date. From thè mid-caudal region on, Wild thought that
sagittal and lateral movements of thè tail were possible.
However, according to Wild, thè overall mobility ot thè
tail did not suggest that it was used for propulsion.
By contrast, Tschanz (1985; 1986; 1988) gave a difter-
ent interpretation of thè morphology described by Wild and
postulated that thè neck of Tanystropheus had little mobil¬
ity. On thè other hand, Tschanz thought that there was rea-
sonable lateral movement in both thè trunk and tail.
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
71
Gastralia
(Pls. I-IV)
Only a few elements are preserved in MSNM BES SC
265 and while thè majority are present in MSNM BES SC
1018, it is difficult to determine their precise number and
arrangement. In thè anteriormost gastral rows two mediai
elements meet along thè midiine. They form an angle,
with thè vertex facing anteriorly. They are thicker medi-
ally and taper laterally into pointed ends. These elements
appear longer than those in thè reconstruction of Wild
(1974: fìg. 36). Additional elements are thinner, curved,
and pointed at both ends. They possibly represent thè lat-
eral elements described by Wild. Wild counted 25 gastral
rows but judging from MSNM BES SC 1018, this number
might have been higher.
archosauromorph-like calcaneal tuber. The astragalus
is an elongate element with an oblique orientation. The
astragalus and calcaneum meet along a straight line. Their
mutuai mobility was precluded, and thè two elements can
be considered as a single unit (Wild, 1974: 118). In some
Tanystropheus specimens, such as MSNM BES SC 265
(Fig. 65) and PIMUZ T 2480 (Fig. 63), thè astragalus has
a proximal lateral process overlapping or indenting with
thè calcaneum. However, Tanystropheus does not show
thè diagnostic features of a complex concave-convex
articulation between thè astragalus and thè calcaneum
(contra Brinkman, 1981: 7).
Dorsally and distally, as previously noted by Wild
(1974) in thè partially articulated tarsus of PIMUZ T
2480 (Fig. 63) and confirmed by MSNM BES SC 1018
Could Tanystropheus walk?
The pes in Tanystropheus
The new specimens offer remarkable new information
on thè morphology of thè pes in Tanystropheus.
The reconstruction of thè tarsus is largely based on
thè perfectly articulated tarsus of MSNM V 3730 (Figs.
35, 61) and thè disarticulated, yet perfectly preserved ele¬
ments in MSNM BES SC 1018 (Fig. 62).
The tarsus of Tanystropheus comprises four ossi-
fied elements. The calcaneum has a polygonal, laterally
rounded shape with thickened lateral margin. Brinkman
(1981: 7) claimed thè presence of a “laterally directed
dorsoventrally compressed tuber on thè calcaneum”
in Tanystropheus. However, there is neither a proces-
sus lateralis ( sensu Schaeffer, 1941: 442) or a lateral,
Fig. 62 - Tanystropheus longobardicus, MSNM BES SC 1018, left pes
in piantar view, particular of thè tarsus. The astragalus and calcaneum
are exposed in dorsal view. I-V = metatarsals. Scale bar 5 mm. Photo:
Massimo Demma.
Fig. 61 - Tanystropheus cf. longobardicus, MSNM V 3730, right tarsus
in piantar view. I-V = metatarsals. Drawing: Massimo Demma.
Fig. 63 - Tanystropheus longobardicus, PIMUZ T 2480, right tarsus in
dorsal view. Scale bar 5 mm. Photo: Heinz Lanz, PIMUZ.
72
STEFANIA NOSOTT1
(Fig. 62), thè astragalus and calcaneum both contrib-
ute to an embayment, in which thè astragalus receives
distai tarsal four, and thè calcaneum receives metatar-
sal V. In MSNM V 3730 (Figs. 35, 61) thè tarsus is
exposed in piantar view (p. 42). Medially, thè astra¬
galus is recessed into a shallow fossa, like in thè left
tarsus of MSNM BES SC 265 (p. 19). However, an
embayment formed by thè astragalus and calcaneum
proximally delimited by a sharp margin cannot be
seen. At thè same time, distai tarsal four and metatarsal
V are not overlapped by thè astragalus and thè calca¬
neum. Consequently, distai tarsal four and metatarsal V
must have possessed rounded proximal articular heads
mirroring thè concavity of thè embayment (Fig. 64).
The presence of a hinge-like mesotarsal joint in Tanys-
tropheus is thus confirmed (Wild, 1974, after Kuhn-
Schnyder, 1960). Wild thought that thè presence of a
mesotarsal joint was one of thè characters indicative of
lepidosaurian affinities. However, thè mesotarsal joint
in Tanystropheus is quite different from that observed
in thè extant squamates, in which thè astragalocalca-
neum and thè distai tarsals have a complex, highly spe-
cialized morphology, permitting rotatory movements
of thè crus on thè pes, as well as flexion-extension
(Rewcastle, 1980). The fourth ossifìed element of thè
tarsus, distai tarsal three, is positioned mediai to distai
tarsal four, between thè astragalus, proximally, and
metatarsal III, distally.
I ascertained that thè specimen of Tanystropheus
recently described by Renesto (2005: pi. 2 H) corre-
sponds to thè description given above with respect to
thè shape of thè tarsal elements and thè nature of their
articulation.
The morphology of thè tarsus in MSNM BES SC 265
(p. 19, Figs. 9, 65), preserved in piantar view, looks dif-
Fig. 64 - Reconstruction of thè tarsus in small-sized specimens of
Tanvstropheus lortgobardicus . A) in dorsal view; B) in piantar view.
Drawing: Massimo Demma.
ferent from MSNM BES SC 1 0 1 8 (Fig. 62) and MSNM V
3730 (Figs. 35, 61), and is more difficult to interpret. The
astragalus appears to have a slightly different shape and
is not obliquely oriented within thè tarsus. The articular
relationships of distai tarsal three and distai tarsal four
with thè astragalus and calcaneum, and between each
other, remain unclear.
Wild (1974: 118) conjectured that thè tarsus of
Tanystropheus originally comprised elements that were
not ossifìed. This cannot be confirmed on thè basis of
thè new material. However, in MSNM BES SC 265 thè
region mediai to thè astragalus is difficult to interpret: it
neither looks like bone nor matrix (Fig. 65, PI. I). It is
possible that it represents a chondrogenic focus or a film
of organic matter, indicative of thè potential for chon-
drifìcation of thè centrale. Indeed, in other protorosaurs,
including Langobardisaurus and Macrocnemus, thè area
distai and/or mediai to thè astragalus is occupied by an
ossifìed centrale (Fig. 66). In MSNM BES SC 265 there
is an undoubted gap mediai and distai to thè astragalus.
The same gap is present in thè perfectly articulated tarsus
of MSNM V 3730. 1 suggest that this gap is indicative of
a cartilaginous element keeping thè metatarsals separate
from thè astragalus and providing some flexibility and
elasticity for thè mediai area of thè tarsus. In some ple-
siosaur limbs there is also indication of thè presence of
elements which apparently do not ossify (Forrest R., pers.
comm., 2007).
In thè well preserved specimens described here,
metatarsals I-IV are tightly packed, their proximal ends
overlapping partly. In MSNM V 3730 (Figs. 35, 61) an
obliquely oriented articular facet is visible on metatarsal
IV, suggesting that metatarsal V overlapped it. If similar
articular facets were present on all thè metatarsals, rela¬
tive movements of thè metatarsals could not be lateral,
as stated by Wild (1974: 118) but must mostly have been
dorso-plantar. In other words, mutuai movements of thè
metatarsals would have resulted in an arched, ventrally
concave metatarsus, rather than in a spreading of thè foot
due to abduction of thè digits.
Fig. 65 - Tanystropheus longobardicus, MSNM BES SC 265, left
tarsus in piantar view. I-V = metatarsals. Scale bar 5 mm. Photo: Mas¬
simo Demma.
TANYSTROPHEVS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
73
C
D
Fig. 66 - A) Langobardisaurus tonelloi (MFSN 1921), left pes in piantar
view, redrawn after Renesto et al., 2002: fig. 5 B. B) Langobardisaurus
pandolfii (MCSNB 2883), left tarsus in piantar view, redrawn after
Renesto, 1994: fig. 7. C) Langobardisaurus pandolfii (MCSNB 2883),
right tarsus in piantar view, redrawn after Renesto, 1994: fig. 8 B.
D) Macrocnemus bassanii (PIMUZ T AIII/208), left tarsus in dorsal
view, redrawn after Rieppel, 1989: fig. 8 f. Not to scale.
The morphology of metatarsal V (in piantar view) is
best seen in MSNM V 3730 (see description on p. 42;
Figs. 35, 61). Metatarsal V is generally described, in
Tanystropheus and its sister-taxa, as “hooked” (Olsen,
1979; Jalil, 1998; Renesto & Avanzini, 2002; Renesto
et al., 2002). The use of this term has been quite sub-
jective over thè years, as recently discussed by Riep¬
pel et al. (2003: 370). “Hooking” of metatarsal V in
Tanystropheus refers only to thè fact that thè roundish
proximal head of metatarsal V articulates both with thè
calcaneum, proximally, and distai tarsal four, medially.
A rounded process lateral to thè distai end of metatar¬
sal V might represent a lateral piantar tubercle for thè
insertion of thè gastrocnemius muscle (Robinson, 1975;
Brinkman, 1980; Rieppel, 1989; Renesto et al., 2002).
In addition, an “outer process” (sensu Robinson, 1975)
with a distinct tuberosity is developed proximo-later-
ally. This might represent thè same process described
by Rieppel (1989) for Macrocnemus, for thè possible
insertion of muscles involved in thè rotation of thè
foot.
The phalangeal formula of thè pes in Tanystropheus
(Tab. 9) is primitive. The first phalanx of digit five takes
thè form of a metatarsal.
The hindlimb: terrestrial versus aquatic locomotion
Once it was ascertained that Tanystropheus was
not a flying reptile, as originally hypothesized by
Bassani (1886) and von Nopcsa (1923), Peyer (1931)
interpreted Tanystropheus as a lizard-like, terrestrial
reptile.
Later, Wild (1974) suggested a terrestrial habit for
thè juveniles but a predominantly aquatic habit for
thè adults. Wild considered thè hindlimb to be thè pri-
mary organ for locomotion in water, mainly because
of a paddle-shaped, webbed foot. According to Wild,
skeletal correlates for aquatic adaptation were thè
wide sites for thè insertion of adductor and retractor
limb musculature on thè ischium and thè femur, thè
elongation of thè metatarsus, and thè presence on thè
metatarsals of oblique articular facets permitting thè
digits to be spread. Wild suggested that digit five, in
particular, had great lateral mobility, and because of
its long, metatarsal-like first phalanx, it contributed
to a large extent to thè distai widening of thè surface
of thè supposed webbed foot. At thè same time, Wild
considered Tanystropheus to be capable of moving
on land as an adult, although he did not explain how
in any detail. In essence, he envisaged a lizard-like
gait, in which thè mesotarsal joint in thè tarsus of
Tanystropheus played a role that is substantially simi-
lar to thè function of thè same joint in extant squa-
mates (Rewcastle, 1980). Following Kuhn-Schnyder
(1960), Wild (1974: 109-110, 142) thought that thè
forelimb was used not only for locomotion on land
but also for digging. Putative skeletal correlates for
this adaptation were an alleged shortened and wide
manus with thickened phalanges and strong claws
(but see p. 33).
Interpretations of Tanystropheus as a fully aquatic
reptile emphasized thè role of thè hindlimb in support-
ing aquatic locomotion (Tschanz, 1985; 1986), or as
thè main organ of propulsion in water (Taylor, 1989).
However, Renesto (2005) recently questioned an
aquatic mode of life for Tanystropheus and discussed
thè morphology of thè limbs. According to Renesto
(2005: 388), thè architecture of thè hindlimb is com-
pletely different from that of any appendage-propelled
aquatic vertebrate and lacks any evident adaptation
for swimming in open waters. He maintained that thè
hindlimb was thè main propulsive force in locomotion
in Tanystropheus, because thè forelimb is relatively
very short and thè carpus poorly ossified. In fact,
forelimbs considerably shorter than thè hindlimbs is
an archosauromorph feature shared by Tanystropheus
with thè relatively well known protorosaurian taxa
Langobardisaurus (Renesto et al., 2002) and Macroc¬
nemus (Rieppel, 1989), and with thè poorly described
Tanytrachelos (Olsen, 1979) and Cosesaurus (Ellen-
berger, 1977; Sanz & López-Martinez, 1984; Peters,
2000a) (Tab. 8).
The new specimens described here confirm that
thè architecture of thè pelvis and hindlimb in Tanys¬
tropheus is very similar to that of thè presumed
terrestrial taxa Macrocnemus (Rieppel, 1989) and
Langobardisaurus (Renesto, 1994; Renesto et al.,
2002). However, major differences, are observed in
thè pes.
74
STEFANIA NOSOTTI
Table 8 - Length ratios between different elements of thè
forelimb and thè hindlimb in thè protorosaurs Tanystro-
pheus longobardicus, Tanytrachelos ahynis, Langobardi-
saurus tonelloi, Cosesaurus aviceps and Macrocnemus
bassanii. The length of thè manus and pes is consid-
ered to be thè length of thè longest unit “metacarpal/
metatarsal+phalanges”.
1) Tanystropheus longobardicus, MSNM BES SC 265.
2) Tanystropheus longobardicus, MSNM BES SC 1018.
3) Tanytrachelos ahynis, YPM 7622, Olsen, 1979.
4) Langobardisaurus tonelloi, MFSN 1921, Muscio G.,
1997 and pers. comm., 2006.
5) Cosesaurus aviceps, MGB-V1, Sanz & López-Mar-
tinez, 1984.
6) Macrocnemus bassanii, MSNM BES SC 111.
The pelvic girdle of Tanystropheus is small compared
to thè size of thè hindlimb and is very similar in shape
and proportions (relative to thè hindlimb) to that of Mac¬
rocnemus (Rieppel, 1989: fig. 4 B). Langobardisaurus
(Renesto et al., 2002: fig. 3 A) has a distinct, posteri-
orly projecting process on thè ischium, which is absent
in Macrocnemus and Tanystropheus. Contra Renesto
(2005) and Rieppel (1989), Tanystropheus does have a
preacetabular process (Fig. 25) similar to that of Lango¬
bardisaurus (Renesto et al., 2002), yet it remains smaller
than that of Macrocnemus . In thè latter, thè preacetabu¬
lar process is robust and tumed outward in an unusual
manner (Rieppel, 1989: 384, fig. 6).
The femur of Tanystropheus (Figs. 1, 8, Pls. I-IV) is
long and stender, and only slightly expanded proximally.
There is no mediai deflection of its proximal articular
head (Fig. 26). This feature indicates a sprawling gait.
The distai end of thè femur displays rounded condyles
(Fig. 29). The tibia and fìbula are of sub-equal length
(Figs. 1, 8, Pls. I-IV, Tab. 5); a spatium interosseum is
present. The tibia has slightly and equally expanded ends,
thè stender fibula has no expanded ends. The proximal
articular surfaces of thè tibia and thè fibula are fiat or
slightly convex. The knee joint (Fig. 29) is apparently
hinge-like and fully functional.
The tarsus of Tanystropheus differs from that of Mac¬
rocnemus and Langobardisaurus (Fig. 66) in having only
four ossified elements (Fig. 61) and a distinct mesotarsal
joint between thè proximal and distai elements (Fig.
64). With thè exception of PIMUZ T 2480 (Fig. 63), on
which Wild based thè reconstruction of thè pes (Wild,
1974: fig. 77), thè elements of thè tarsus in thè Tanystro¬
pheus material in thè PIMUZ Collections are invariably
displaced (Wild, 1974: figs. 74-76). This might suggest a
looser articulation than that observed in thè very compact
tarsus of Macrocnemus and Langobardisaurus . A dislo¬
cation of thè crus and tarsal ossifications seems to be thè
generai rule. It is indicative of a looser crurotarsal joint
in Tanystropheus, than in Macrocnemus (Rieppel, 1989:
fig. 8; Fig. 66 D in this paper) and Langobardisaurus
(Fig. 66 A-C), although overall thè ankle joint is quite
similar in all three taxa. In Langobardisaurus thè tibia
and thè fibula fit loosely into concavities formed by thè
proximal borders of thè astragalus, calcaneum and cen¬
trale (Renesto et al., 2002). In Macrocnemus (Rieppel,
1989), thè fibula is received in an embayment formed
by thè astragalus and calcaneum. The tibia articulates
with thè astragalus, hearing a distinct articular facet on
its mediai side. This facet forms thè proximal part of an
embayment completed by thè centrale and distai tarsal
one, within which thè tibia is accomodated during thè
propulsive phase ofthe stride (Rieppel, 1989). In Tanys¬
tropheus, thè articular facet formed by thè astragalus
and calcaneum for thè fibula is similar to that described
in Macrocnemus. There is no identifiable facet for thè
tibia on thè astragalus. However, in MSNM BES SC 265
(Fig. 65) thè tibia, even though it is displaced medially,
stili contacts thè astragalus. The presence of a carti-
laginous centrale in thè tarsus of Tanystropheus, as dis-
cussed above, is consistent with thè configuration of thè
ankle joint in other protorosaurs, in which a contact of
thè tibia with thè centrale is observed (Benton & Alien,
1997; Dilkes, 1998).
Dislocation of thè tarsus and metatarsus is commonly
observed in thè fossil material of Tanystropheus. By con-
trast, thè metatarsals are in most cases preserved in asso-
ciation, with their proximal ends overlapping. As in other
protorosaurs, this suggests that thè metatarsals formed a
functional unit. Metatarsal V has an outer process and a
lateral tubercle similar to thè presumed terrestrial Macroc¬
nemus (Rieppel, 1989) and Langobardisaurus (Renesto et
al., 2002). The presence in thè foot of Tanystropheus of
a metatarsal-like first phalanx on digit five is a character
shared with other protorosaurs, in particular Tanytrach¬
elos, Langobardisaurus and Cosesaurus. In Tanystroph¬
eus, however, thè articulated phalanx exceeds thè length
of metatarsals I-IV, while in Tanytrachelos, Lango¬
bardisaurus and Cosesaurus it is shorter than thè longest
metatarsal (metatarsal III in Tanytrachelos, Olsen, 1979:
fig. 4 A; metatarsal IV in Langobardisaurus, Renesto et
al., 2002: fig. 5, and Cosesaurus, Ellenberger, 1977: fig.
12). The phalangeal formula of thè pes in Tanystropheus
(2, 3, 4, 5, 4) is thè same as in Tanytrachelos, Langobardis¬
aurus and Macrocnemus (Tab. 9).
Comparing thè limbs of Tanystropheus with those ot
other protorosaurs, and, in particular, those ofthe recently
discovered Chinese protorosaur Dinocephalosaurus (Li,
2003; Li et al., 2004), Renesto (2005) concluded that
thè morphology of thè limbs of Tanystropheus rules
out any kind of aquatic locomotion. He also empha-
sized that none of thè authors who had proposed a fully
aquatic mode of fife for Tanystropheus outlined which
mode of propulsion was adopted by this reptile. In fact,
Tschanz (1985; 1986) did suggest that Tanystropheus
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
75
Table 9 - Comparison of thè phalangeal formulae of thè manus and pes in thè protorosaurs Tanystropheus
longobardicus, Tanytrachelos ahynis, Langobardisaurus , Cosesaurus aviceps, and Macrocnemus bassanii.
was an axial-subundulational swimmer ( sensu Braun &
Reif, 1982; 1985), in which locomotion resulted from
undulatory movements of thè trunk and tail, addition-
ally supported by paddling of thè hindlimbs. However,
Tschanz did not discuss in detail thè morphology of thè
hindlimb in Tanystropheus and did not explain how this
paddling was possible.
Assuming an aquatic mode of life for Tanystropheus ,
it is stili most likely that it retumed to land for reproduc¬
tion, in which case it must have been capable of moving
on land in some way. However, nobody has ever tried
to explain in detail how this reptile walked. Although a
detailed analysis of locomotion is beyond thè scope of
this paper, some points are raised below.
The absence of articular cartilages in thè fossil mate¬
rial limits a complete understanding of thè full range of
movements possible at any joint. The shape of thè femur
apparently indicates a sprawling gait for Tanystropheus.
Rieppel (1989) pointed out that with a sprawling gait
there is a requirement for some rotation of thè lower
limb relative to thè femur, and that in extant squamates
this rotation does not take place within thè knee joint, as
demonstrated by Rewcastle (1980) but in thè mesotarsal
joint. In Macrocnemus , in thè absence of such a joint, as
well as of an archosauromorph-like articulation between
thè astragalus and calcaneum, Rieppel (1989) stated
that thè rotation took place between thè crus and thè
proximal tarsal ossifications (p. 74, Fig. 67). As a result
of this movement, thè distai metatarsal heads would be
aligned at right angles to thè body axis, thereby distribut-
ing thè body weight during thè propulsive phase of thè
stride. Renesto et al. (2002) assumed a similar rotation
movement in Langobardisaurus. In thè latter, however,
thè presence of a larger and more transversally elongate
centrale, thè lack of distai tarsal one, and thè presence of
a second axis of rotation between thè centrale-astragalus-
calcaneum and thè distai tarsals-metatarsals, would have
resulted in a more anterior orientation of thè pes than in
Macrocnemus (Renesto et al., 2002).
Ongoing research indicates a digitigrade stance in
thè pes of many protorosaurs, including Tanytrachelos
(Peters, 2000b: figs. 9 B, 15 A), Cosesaurus (Peters,
2000b: fig. 16), Macrocnemus (Avanzini & Renesto,
2002) and Langobardisaurus (Renesto et al., 2002), and
to a bipedal posture, during rapid locomotion - as previ-
ously stated by Rieppel (1989) for Macrocnemus - or even
while standing and walking.
For Tanystropheus, a digitigrade stance in thè pes is
thè only plausible configuration while walking as well.
Any analysis of terrestrial locomotion in Tanystropheus
must take into account thè metatarsal-like proportions
of thè first phalanx of digit five during pedal plantar-
flexion {sensu Brinkman, 1980: 278). As mentioned
above, thè length of this phalanx, when articulated with
metatarsal V, exceeds that of thè longest (III) metatarsal.
Intuitively, it is clear that only thè elements of thè pes
distai to thè distai end of thè metatarsal-like phalanx
could contact thè ground. Furthermore, thè metatarsus
of Tanystropheus is asymmetrical, although not in thè
same manner as Macrocnemus, Langobardisaurus and
Cosesaurus. In thè latter taxa there is a regular increase
in length from metatarsal I through metatarsal IV. In
Tanystropheus, thè longest metatarsal is metatarsal III,
followed by metatarsal IV, metatarsal II and metatarsal
I. Tanytrachelos (Olsen, 1979: fig. 4) apparently has a
similar metatarsal configuration. Again, however, thè
length of thè articulated metatarsal-like phalanx of digit
five in Tanytrachelos does not exceed thè length of thè
metatarsals. Analysing thè hinge lines (Peters, 2000b)
and comparing thè pes of Tanytrachelos with thè Gwyn-
nedichnium trackmaker, Peters (2000b: figs. 9 B, 15 A)
suggested that thè pes of Tanytrachelos could adopt both
a plantigrade and digitigrade configuration. Peters also
drew hinge lines in thè pes of Tanystropheus, as recon-
structed by Wild, briefly discussing thè pattern obtained
(Peters, 2000b: 30, fig. 15 B). Nevertheless, he was
unable to come to a definitive conclusion about thè pos¬
sible configuration for thè pes in this protorosaur. A pre-
liminary examination of thè hinge lines on thè superbly
preserved left pes of MSNM BES SC 1018 shows a
slightly different pattern than that figured by Peters in
thè plantigrade configuration. The mediai and transver¬
sai set of lines are more distai (Ma, Ta and Tb in Peters’
fig. 15 A were not identified in thè pes of MSNM BES
SC 1018), and clearly distai to thè articulation of thè
metatarsal-like phalanx of digit five and thè subsequent
phalanx. The whole lateral set of lines is too tangential
to have a functional meaning.
Any configuration of thè pes in Tanystropheus requires
its re-orientation to bring a set of hinge lines oriented at
right angles to thè direction of locomotion. While thè
ankle joint in Tanystropheus is similar to that of Macroc¬
nemus and Langobardisaurus , in thè absence of an ossi-
fied centrale, if any, in Tanystropheus, there is no solid
76
STEFANIA NOSOTTI
embayment constraining thè dislocation of thè tibia along
thè medio-distal margin of thè tarsus. This dislocation is
responsible for rotation and consequently thè outward ori-
entation of thè pes. The main movement permitted at thè
crurotarsal joint in Tanystropheus was probably flexion
and extension of thè ankle.
In contrast to Macrocnemus and Langobardisaurus,
a (poorly specialized) mesotarsal joint is present in thè
tarsus of Tanystropheus . In thè reconstruction presented
here (Fig. 64, p. 72), such a joint would have permitted
thè dorsiflexion of thè unit comprising thè distai tarsals
and metatarsals on thè unit comprising thè astragalus
and calcaneum. A little mutuai rotation of thè two units
might have been possible but I doubt that thè extent of
this movement permitted thè re-orientation of thè pes
required for thè propulsive phase of thè stride.
On thè other hand, thè prevalent movement of flex-
ion-extension at thè ankle and at thè mesotarsal joint in
Tanystropheus , together with thè unossified mediai part of
thè tarsus and thè loose joints between thè tarsal elements
- and also between thè tarsus and thè crus - are consistent
with a paddling motion. The overall flexibility of thè pes
was apparently more important than thè mobility at each
joint.
Skeletal correlates for an efficient pedal plantar-
flexion (Rieppel, 1989) include a well differentiated
“hooked” (p. 73) metatarsal V, with an outer process
and distinct lateral and mediai tubercles, and elongate
metatarsals I-IV forming a single functional unit. The
occurrence of some of these features in Tanystropheus
might be interpreted as plesiomorphic and inherited
from a common ancestry within thè terrestrial proto-
rosaurs.
Tanystropheus mode of life: on what side of thè
shoreline?
The skeletal anatomy of Tanystropheus is unique and
there is no equivalent in either extant or extinct animals.
The recently described Chinese protorosaur Dinocepha-
losaurus (Li, 2003; Li et al., 2004) has an overall form
closely recalling Tanystropheus. However, in Dinocepha-
losaurus thè proportions of thè limbs, with shortened
epipodials, thè poorly ossified carpus and tarsus, thè
rounded astragalus and calcaneum, and thè poor differen-
tiation of metatarsal V, unequivocally indicate thè adapta-
tion to an aquatic mode of life.
Criticai issues in thè interpretation of thè mode of life
for Tanystropheus include thè generai proportions and
shape of thè body, skeletal anatomy, mobility of thè long
neck and static problems related to it, locomotion (see
also thè preceding section), and feeding.
Tanystropheus does not have a compact and stream-
lined body (Taylor, 1989). The trunk is very short com-
pared with thè extremely long neck and thè tail. The latter
is neither deep and/or laterally compressed. The gracile
limbs are not paddle-shaped, and neither shortened nor
broadened. The pes is very long relative to thè crus and thè
femur, but similar proportions are observed in thè related,
presumed terrestrial taxa Langobardisaurus and Macroc¬
nemus (Tab. 8). Except for thè disproportionately elongate
neck, thè overall form of thè skeleton of Tanystropheus is
not strikingly different from that of thè latter taxa.
Nevertheless, Rieppel (1989) underlined many
skeletal correlates indicative of terrestrial habits for
Macrocnemus which are absent in Tanystropheus, and
emphasized that thè overall degree of ossification is less
in adult Tanystropheus than in Macrocnemus. This might
well indicate that Tanystropheus was an aquatic animai
throughout its life. In fact, thè mild degree of skeletal
paedomorphosis observed in Tanystropheus might have
important functional effects. As discussed in thè preced¬
ing section, thè morphology of thè pes in Tanystropheus
does not suggest a marked adaptation to an aquatic mode
of life, yet it renders it difficult to imagine efficient ter¬
restrial locomotion. Moreover, thè minimal degree of
ossification of thè carpus and thè reduced size of thè
forelimb suggest that thè forelimb was not a major con-
tribution to any kind of locomotion (Renesto, 2005).
Renesto (2005) emphasized that thè extremely long,
yet scarcely flexible, neck of Tanystropheus seems unsuit-
able in any environment, and summarized an overall view
of previous studies (Renesto, 2005: 387, fig. 10). Accord-
ing to Peyer (1931), Wild (1974) and Kummer ( 1 975) thè
neck of Tanystropheus was rather mobile and held hori-
zontally or considerably raised.
The axial locomotory System in Tanystropheus was
more recently interpreted by Tschanz (1985; 1986) draw-
ing a comparison with extant reptiles (Iguana and Vara-
nus), and thè neck was concluded to have been almost
inflexible. Tschanz compared thè neck of Tanystropheus
to an architectural construction in which thè interspinal
ligaments and thè intervertebral muscles represent a ten-
sion boom between two neighbouring vertebrae, while thè
bundles of cervical ribs represent a strut. The final result
would be a neck incapable of dorsal flexion. Tschanz
maintained that thè neck could only be held horizontally
and not raised above thè level of thè shoulder. According
to Tschanz, this configuration is strongly indicative of a
fully aquatic mode of life for Tanystropheus. Buoyancy
would have supported thè long neck in thè water and
would have reduced thè hydrostatic stress on thè cardio-
vascular System.
By contrast, according to thè most recent interpreta¬
tion by Renesto (2005), there would be evidence indicat-
ing that thè neck was more flexible and mobile than sug-
gested by Tschanz. The shafts of thè cervical ribs, running
in tight bundles ventral to thè vertebral column, would
have at least partially constrained thè dorsoventral flex¬
ion of thè neck but there would have been some limited
flexibility, so that thè neck was not completely rigid. As
a result of thè hollow vertebral centra, it was very light
and, even in thè absence of a strong epaxial musculature,
it could have been supported on land, held in an inclined
posture. Drawing a comparison with thè long-necked
azdarchid pterosaurs, Renesto (2005) conjectured that
Tanystropheus likewise possessed a muscle/ligament
System allowing thè neck to be raised above thè horizon-
tal piane.
According to Renesto (2005), thè form of thè 12*
cervical vertebra in Tanystropheus would account for thè
inclined posture of thè neck. Its forward-slanting poste-
rior surface, when in juxtaposition with thè fiat anterior
surface of thè fìrst dorsal vertebra, would produce an
angle at thè base of thè neck. However, thè presence of
an asymmetrical intervertebral disc (Tschanz, 1986: 65)
or a slightly dorsally curved vertebral column in thè trunk
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
77
region might well result in a horizontally stretched neck,
even in thè presence of thè forward-slanting posterior sur-
face of thè 12th cervical vertebra.
Conceming thè static problems related to thè presence
of a long neck in thè terrestrial environment, Renesto
(2005) emphasized that thè whole anterior portion of thè
body of Tanystropheus would have been lighter than thè
posterior one, and speculated that a large muscular mass
at thè base of thè tail shifted thè centre of mass posteri-
orly, thus stabilizing thè body. He cited as evidence for
thè presence of this muscular mass thè massive patches
of black matter and a large number of small spherules
comprising tiny calcium carbonate crystals at thè base
of thè tail. It is not clear, however, whether this type of
preservation is related to peculiar taphonomical condi-
tions characterizing thè levels of thè Lower Meride
Limestone from which thè specimen - collected from an
isolated block - is alleged to have come (Renesto, 2005:
378), and whether it is observed in other fossil vertebrates
from thè same levels. This would support Renesto’s inter-
pretation. I concur with Renesto that thè presence of thè
long pleurapophyses of thè proximal caudal vertebrae is
related to muscular insertions. At thè same time, many
extant reptiles, including lizards, have caudal lipid Stores.
In thè viviparous skink Niveoscincus metallicus caudal fat
bodies comprise thè majority (55-78%) of thè fat reserves,
and thè 90-95% of caudal fat occurs within thè proximal
third of thè tail (Chapple & Swain, 2002). Finally, stabi-
lization of thè body due to thè weight of a posterior, large
muscular mass might also be equally advantageous in an
aquatic animai.
Analysing thè patterns of aquatic locomotion within
thè Sauropterygia and discussing thè probable steps in
thè origin of aquatic locomotion among reptiles, Car¬
roll & Gaskill (1985) pointed out that in thè early stages
of aquatic adaptation propulsion in water is mainly
achieved through lateral undulations of thè axial skel¬
eton. In thè majority of secondarily aquatic reptiles,
use of limbs is disadvantageous in water because of thè
constraints imposed by thè morphology and stereotyped
pattern of movement inherited from dose terrestrial
ancestors. Therefore, thè emphasis put on thè forelimbs
for propulsion that is observed in nothosaurs and plesio-
saurs can be interpreted as secondary, and subsequent to
a progressive reduction of thè forelimbs to limit drag in
thè early stages of aquatic adaptation. Extant crocodiles
and thè marine iguana also swim with lateral undula¬
tions of thè trunk and tail, keeping thè forelimbs dose
to thè body and mostly trailing thè hindlimbs, which
are occasionally thrust backwards in a paddling motion
(Carroll & Gaskill, 1985). On thè other hand, Braun
& Reif (1985) stated that discontinous propulsion,
because of its low effìciency at medium to high Rey-
nolds-numbers, plays practically no role in fish. At thè
same time, discontinuous propulsion such as paddling
occurs in semi-aquatic tetrapods which can also walk,
ri fly, jump, and even climb. Usually, adaptation to rowing
and paddling requires minor anatomical changes of thè
| terrestrial locomotory apparatus (Braun & Reif, 1985).
In thè light of thè discussion above, Tschanz’s (1985;
1986) interpretation of Tanystropheus as an axial-subun-
dulational swimmer ( sensu Braun & Reif, 1982; 1985),
which might have also used thè hindlimbs as paddles,
seems consistent.
nV
Other long-necked reptiles, such as thè plesiosauroid
plesiosaurs, have been interpreted as relatively slow
swimmers living in Coastal environments. While they had
evolved efficient propulsive mechanisms, plesiosaurs, in
particular long-necked plesiosauroids, have a poor hydro-
dynamic profile (Massare, 1988): thè long neck, increases
drag. In thè diagram of “fineness ratios” published by Mas¬
sare (1988: fìg. 2), some plesiosauroids do not fall into thè
category representing body shapes which minimize thè total
drag and which are optimal for fast, continuous swimmers.
According to Massare, many of thè plesiosauroids had a
slower cruising swimming speed for a given size than fusi-
form-shaped marine reptiles such as ichthyosaurs and plio-
sauroids. Forrest R. (pers. comm., 2007) emphasizes that,
from a hydrodynamic point of view, a stiff neck is essential
and there is a growing consensus that thè neck flexibility in
some plesiosauroids was akin to a stiff fishing rod (see also
Ford, 2002). Interpreting Tanystropheus as an axial-subun-
dulational swimmer, Tschanz (1985; 1986) underlined that
in this kind of swimmer thè anterior part of thè body must
be rigid, and that thè stiffened part of thè body in Tanystro¬
pheus is represented by thè elongate neck.
The new Tanystropheus specimens confimi that thè
articular surfaces of thè pre- and postzygapophyses of thè
dorsal vertebrae posterior to thè second-third and of thè
caudal vertebrae are oriented more or less horizontally,
and therefore permit lateral movements. While Tschanz
(1985) stated that thè caudal haemapophyses are mod-
erately elongate, I concur with Renesto (2005) in saying
that, on thè basis of skeletal evidence, thè tail in Tanystro¬
pheus is not powerful, enlarged and laterally flattened as it
would be expected in a tail-propelled tetrapod. However,
albeit highly speculative, thè hypothesis that thè shape
of thè tail in thè living animai was different from that
indicated by thè skeletal morphology cannot be excluded.
Thus, for istance, it is possible that fins composed of
soft tissue were present in thè living animai. According
to Renesto (2005), thè elongate pleurapophyses of thè
proximal caudal vertebrae would have hindered lateral
undulation of thè tail. However, thè pleurapophyses in
specimen MSNM BES SC 265 (Fig. 6) are not too long to
completely preclude lateral movement, and, if thè length
of thè tail is taken into account, little lateral movements at
any vertebral joint might well have produced an appreci-
able lateral movement of thè tail as a whole.
In thè light of thè discussion above, I regard Tanys¬
tropheus as an aquatic protorosaur with dose terrestrial
ancestors, living in shallow waters (maximum a few ten
meters deep, Rhòl et al., 2001) and in all probability
retuming to land for reproduction. I concur with Tschanz
(1985; 1986) that Tanystropheus was a slow, not highly
specialized swimmer, relying on lateral undulations of thè
trunk and tail for aquatic propulsion, perhaps enhanced by
paddling with thè hindlimbs.
A criticai, stili unclear issue in thè interpretation of thè
mode of life for Tanystropheus is feeding, in particular
method of capture and processing prey. Stomach contents
were reported by Wild (1974: 51, 142) in some large-
sized specimens of Tanystropheus. They consist of fish
and hooklets from cephalopods’ arms. This suggests that
Tanystropheus had thè capability to seize elusive prey.
However, it is difficult to imagine how it could catch such
preys both in water and from thè shoreline, especially if
thè neck was stiff.
78
STEFANIA NOSOTTI
As discusseci by Massare (1988), thè swimming capa-
bilities of Mesozoic marine reptiles have implications for
thè mode of predation. Plesiosauroid plesiosaurs provide
a model of feeding strategies that might be adopted by
an aquatic, long-necked animai. Because of their slower
continuous swimming speeds, plesiosauroids had thè
option for one of three strategies: pursue slow prey, eat
sessile prey or use an ambush technique to capture prey
(Massare, 1988). According to Massare, thè possibility of
an ambush mode of attack is more speculative for thè ple¬
siosauroids than for thè crocodiles and mosasaurs. How-
ever, very long-necked plesiosauroids might have been
ambush predators, thè prey being caught before thè large
body was even detected in dark, murky water (Massare,
1988). There is also evidence that thè four flipper “gait”
of plesiosaurs, while not especially efficient for normal
locomotion, was very well suited for rapid acceleration
required in thè ambush method of predation (Long Jr. et
al., 2006). McHenry et al. (2005) recently found stom-
ach contents dominated by benthic invertebrates in two
Australian elasmosaurid specimens, suggesting that even
structures as specialized as thè elasmosaurid neck are not
necessarily indicative of very specialized niches. This is
confìrmed by thè wide range of tooth forms found in thè
long-necked plesiosaurs (Forrest R., pers. comm., 2007).
Contra Wild (1974: 142), it seems very unlikely that
an aquatic Tanystropheus was a fast swimmer pursuing
slower prey. An ambush method of capturing prey, requir-
ing fast starts and rapid acceleration of thè body, i.e. short
bursts of swimming, also seems improbable for Tanystro¬
pheus (but see Ford, 2002).
An alternative ambush method of capturing prey was
hypothesized by Peters (2005) for thè recently described,
aquatic protorosaur Dinocephalosaurus. According to
Peters, this animai was a sit-and-wait predator, hiding in
bottom silt and snatching passing fish from below. Peters
considered thè same model applicable to Tanystropheus as
well. Li et al. (2004) argued that extending thè head verti-
cally would have been impossible for Dinocephalosaurus
because thè hydrostatic pressure would have prevented
lung inflation. Peters (2005) suggested that this problem
could have been overcome by gulping a small bubble of
air and carrying it to thè bottom in thè throat sac before
passing it to thè lungs under equalized pressure. Contra
Peters, LaBarbera & Rieppel (2005) maintained that
Dinocephalosaurus was unlikely to have been a benthic
ambush predator, because its orbits are not facing dorsally
and thè morphology of thè cervical vertebrae indicate that
dorsiflexion of thè neck was impossible. Moreover, such
motion would generate high drag forces on thè neck that
would tend to drag thè body of thè animai in thè direction
opposite to thè motion of thè head.
Li et al. (2004) maintained that a form of suction
feeding was a possible strategy for Dinocephalosaurus.
Moderate lateral flexion of thè neck, followed by rapid
straightening and splaying outward ot thè ribs, would
produce an increase in thè esophageal volume, that in
turn would create suction. This model was questioned by
Peters (2005) and Demes & Krause (2005) who denied
thè possibility of an expansion of thè esophagous as
suggested by Li et al. (2004). They also argued that thè
feeding mechanism invoked by thè latter authors would
require physiologic adaptations for removai of salt from
thè swallowed water and/or some mechanism of second-
ary water expulsion. However, LaBarbera & Rieppel
(2005) objected to thè arguments of Peters (2005) and
Demes & Krause (2005), noting that reptilian physiol-
ogy automatically implies thè presence of some form
of salt gland. Interestingly, fossil evidence of salt glands
was recently reported by Femàndez & Gasparini (2000)
for thè marine metrorhynchid crocodiliform Geosaurus.
LaBarbera & Rieppel also suggested that excess water
might have been expelled through thè orifices between
thè fang-like grasping teeth of Dinocephalosaurus .
Taylor (1989) emphasized that thè stiffened neck of
Tanystropheus seems inappropriate for an ambush strat-
egy of prey capture, and suggested that thè residuai lateral
flexibility of thè neck might have been used for lunging
thè head over thè prey. Tschanz also (1985; 1986) sug¬
gested that lunging thè head over thè prey might have
been thè feeding strategy of Tanystropheus, following a
model of “kinetic inerbai feeding”. Given thè extremely
elongate neck, a minor dorsal bending in its posterior part
would have permitted a sufficient retraction of thè head.
The energy stored by thè bent cervical ribs could then
be used to accelerate thè head and manipulate thè prey
deeper into thè pharynx.
Because of thè stiffened neck, it is difficult to imagine
Tanystropheus feeding on fast moving prey. Both a model
of kinetic inerbai feeding ( sensu Tschanz, 1985; 1986) or
suction feeding {sensu Li et al., 2004) implies thè capa-
bility of some flexion of thè neck. The model suggested
by Tschanz seems more plausible, given thè sub-vertical
orientation of thè pre- postzygapophyseal articulation of
thè last cervical and first dorsal vertebrae. This indicates
that thè neck could be moved in thè sagittal piane, while
its lateral movement could be achieved only with a lateral
movement of thè trunk. Because of thè poor knowledge of
thè cranio-cervical joint, movements of thè head relative
to thè neck cannot be estimated.
A functional interpretation of thè dentition might be
relevant for our understanding of thè feeding strategy
adopted by Tanystropheus. Wild (1974: 50-51) compared
thè dentition in thè large-sized specimens of Tanystroph¬
eus (his “adult” individuate) with that of marine predators
such as thè nothosaurs. Ford (2002) first observed that thè
premaxillary and anterior dentary teeth of Tanystropheus
are interlocking and suggested that in this aspect - and in
dorsally placed nares - thè skull of Tanystropheus resem-
bles those of fish-eating pterosaurs and sauropterygians
rather than terrestrial diapsid. Indeed, thè anterior conical
interlocking teeth of Tanystropheus seem well-suited for
catching prey.
The posterior tricuspid dentition of small-sized Tanys¬
tropheus specimens was interpreted by Wild (1974: 50) as
evidence for predominant insectivorous habits. According
to Wild, thè tricuspid teeth were an adaptation for grip-
ping small and fast moving prey, catched with thè long,
conical anterior teeth. The shagreen-like pterygoid teeth
and thè pointed vomerine and palatine teeth confìrmed thè
functional interpretation of thè dentition of Tanystropheus j
as a device for capturing and crushing prey.
Based on thè comparison with thè tricuspid dentition
of thè extant marine iguana, Cox (1985) altematively con-
jectured an herbivorous diet for Tanystropheus. However,
as pointed out by Taylor (1989), thè absence of a very
mobile neck would make it difficult for Tanystropheus to
hunt for insects or tear algae from thè rocks.
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
79
A new model for thè interpretation of thè tricuspid
dentition in Tanystropheus is provided by thè extant pin-
nipeds. These mammals tipycally display cheek teeth (or
postcanines) little differentiated from one another which,
particularly in thè Phocidae, display a considerable vari-
ability in shape and degree of cusp development (Miya-
zaki, 2002a). In generai, cusps are related to gripping of
slippery prey. Tricuspid teeth are observed in many spe-
cies. The species of thè genus Pusa feed on fìsh ( P. sibir-
icà), fish and crustaceans (P. caspica ), or on small fìsh and
a wide variety of small pelagic amphipods, euphausiids,
and other crustaceans (P. hispida ) (Miyazaky, 2002b).
The Harbor Seal ( Phoca vitulina ) feeds opportunistically
on many kinds of fishes, shellfishes and squids (Schef-
fer, 1969). In some species, such as thè Crabeater Seal
( Lobodon carcinophaga), thè cheek teeth have a highly
modified shape with complex elongate cusps to trap and
strain krill (Miyazaki, 2002a).
In a fully aquatic Tanystropheus thè posterior tricus¬
pid teeth might have well been an adaptation to gripping
and piercing slippery, tough-skinned prey, and possibly
cutting flesh. Wild (1974: 50) observed thè similarity
of thè tricuspid teeth of Tanystropheus with those of thè
primitive flesh-eating cynodonts, but argued that they
had not a shearing function. He emphasized that thè tri¬
cuspid teeth in thè PIMUZ specimens show horizontal
rather than oblique wear facets, suggesting a tooth to
tooth occlusion. However, oblique wear facets can be
seen in some maxillary teeth in MSNM BES SC 265.
Moreover, based on thè new specimens from Besano,
thè tricuspid teeth of thè upper jaw are positioned labial
to those of thè lower jaw. The upper and lower set might
perhaps create a shear pattern similar to thè camassial
shear of Carnivora.
Finally, some authors claimed that an interpreta¬
tion of thè mode of life for Tanystropheus based on thè
understanding of thè function of its long neck, might be
misleading (Taylor, 1989). Given thè positive allometric
growth of thè neck in Tanystropheus (Tschanz, 1988;
Wild, 1974), Taylor (1989) considered thè great length
of thè neck might be an indirect consequence of thè
evolution of large size. In this case thè large size might
be an adaptation for a reason other than feeding, such
as sexual competition or defence. Taylor concluded that
perhaps Tanystropheus had a long neck simply because it
was a big animai, and it survived in spite of it rather than
because of it.
Tschanz (1988: 1002) also stated that “structures with
no recognizable adaptive value may be more reasonably
explained as results of allometric growth”. Analysing thè
ontogenetic development of thè neck in T. longobardicus
and Macrocnemus bassanii, Tschanz observed that thè
positive allometric growth of thè neck has different
growth parameters for thè two taxa. In particular, there
would be evidence for a relatively longer neck and for
decelerated growth of it in Tanystropheus , in comparison
to Macrocnemus. According to Tschanz, this would make
sense assuming thè occurrence in thè ancestry of Tanys¬
tropheus, morphologically exemplifìed by M. bassanii, of
heterochronic processes, such as hypermorphosis and pre-
displacement, accounting for an extreme elongation of thè
neck with increasing body size. In thè absence of decel¬
erated growth of thè neck, thè accelerated body growth
would have produced an elongation of thè neck rendering
it a functionally inappropriate or unadapted structure.
Discussing thè elongation of thè neck in different Tanys¬
tropheus species, Wild (1987) apparently maintained as
well that an extremely long neck was unadapted, when
recognizing thè specialization of thè cervical vertebral
region as one of several “evolutionary trends” character-
izing thè history of Tanystropheus and potentially leading
to extinction.
The hypothesis that some structures characterized by
unexplained but demonstrated positive allometric growth
developed simply as a consequence of a positive selective
pressure for a large body size was often put forward to
explain “odd” morphology. As in thè emblematic example
of thè huge antlers of thè Irish Elk, peculiar anatomical
traits were traditionally considered as inadaptive. Dis¬
cussing thè origin and function of “bizarre” structures,
and, in particular, thè case of thè Irish Elk, Gould (1974)
sharply commented on thè interpretation maintaining that
thè enormous antlers are a passive consequence of selec-
tion for larger bodies as follows: “Curiously, this standard
contention has not escaped thè dead hand of thè orthoge-
netic explanation that it bravely claimed to replace. For
it assumes that thè antlers were disadvantageous per se,
and that selection preserved them only because it favored
thè total phenotype of larger bodies and antlers. Instead
of an immutable trend, we now have an immutable cor-
relation. Thus, thè assumption of deleterious antlers was
transported bodily from thè orthogenetic to thè allometric
argument...”. Gould argued that in thè case of thè Irish
Elk, thè “allometric antidote to orthogenesis” rested more
upon dogmatic assumptions than “upon a firm foundation
of careful and copious data”, and cited evidence for a
primary selective advantage for thè antlers, as structures
for display conditioning thè female choice or establishing
dominance through ritualized encounters between com-
peting males.
Zamik (1925) strongly claimed an adaptive meaning
of thè long necks of plesiosaurs: “One has to exclude thè
notion that these animals were thè product of accidental
orthogenesis which we will cali hypermorphology; hyper-
morphological disharmony between form and function
would led to their extinction (as for example, thè ammo-
nites with untwined spirai in thè Cretaceous Period)”.
Recently, Noè (2006) emphasized that thè possession of
a long neck in thè marine environment poses a number of
functional, biomechanical, ecological and physiological
problems. Nevertheless, its continued presence through-
out thè long evolutionary story of thè plesiosaurs dem-
onstrates that a long neck can be a successful adaptation
to life in water. It’s also worth noting that thè tendency to
hyperelongation of necks occurred in at least two, possi¬
bly three distinct evolutionary lineages in plesiosaur evo¬
lution (Forrest R., pers. comm., 2007; O’Keefe & Robin,
2001; Smith, 2003).
Finally, elongation of thè neck is generally interpreted
as an adaptation related to feeding strategies. In fact,
“unhortodox” hypotheses on thè functional meaning of
neck elongation were put forward for some long-necked
animals. This is thè case for neck elongation in extinct
sauropods (Senter, 2007) and in extant giraffe (Sim-
mons & Scheepers, 1996), which traditionally have been
thought to have evolved under thè pressure of competition
for food, but have altematively been indicated as driven
by sexual selection.
80
STEFANIA NOSOTTI
IMPLICATIONS FOR PHYLOGENETIC ANALYSIS
Rieppel et al. (2003) recently presented a synthesis of
thè most recent/computer supported cladistic analyses of
protorosaurian interrelationships, obtained by combining
thè data published by Benton & Alien (1997), Jalil (1997)
and Dilkes (1998). Major difficulties affecting thè results
of this analysis centre on a very poor knowledge of many
protorosaurian taxa, as well as thè description and coding
of thè characters included in thè analysis. These problems
are often related to objectivity in assessing large list of
characters, within a large list of taxa, and with direct
observation of all thè originai material.
It should probably be accepted that some characters
will remain only provisionally phylogenetically informa¬
tive. However, thè importance of first hand systematic
anatomical work of thè originai material cannot be over
emphasized. The new specimens described here confimi
this, providing new information on some poorly defined
or previously undescribed characters of Tanystropheus.
The discussion given in this paper also emphasizes how
thè description of some characters is (inevitably) based
more on subjective interpretation of thè material, rather
than on strong evidence offered by thè material itself.
The characters coded by Benton & Alien (1997;
BA=characters by Benton & Alien), Jalil (1997;
J=characters by Jalil) and Dilkes (1998; D=characters by
Dilkes) are re-considered here on thè basis of thè descrip-
tions and discussions presented in this paper. Contrary
to Benton & Alien, Jalil did not differentiate between T.
longobardicus and T. meridensis but used thè genus as
terminal taxon, while Dilkes considered T. longobardicus
only. I assigne characters States for T. meridensis on thè
basis of preliminary personal observation, provisionally
maintaining it as a separate species. However, thè holo-
type, and thè single known specimen, of T. meridensis
preserves only thè skull and thè first six cervical verte -
brae. Overlapping characters used by Benton & Alien,
Jalil and Dilkes are reported in parentheses. If not men-
tioned, overlapping characters have been coded thè same
by all authors. All thè characters of thè different authors
are provisionally considered but an important next step is
to exclude characters that might be more easily affected
by inaccurate scoring (see also discussion in Rieppel et
al., 2003). These comprise ill-defined characters, and par-
ticularly those including qualitative terms, that are there-
fore highly subjective. Characters that may be influenced
by size, ontogenetic stage, individuai variation or vagar-
ies of preservation also should be re-evaluated. They are
particularly relevant to ongoing discussions conceming
thè different species of Tanystropheus (pp. 5-6).
The coding for thè following characters of thè skull is
confìrmed or emended. Benton & Allen’s coding for BA4,
BA7, BAIO, BAI 2 (J43, D35), and BAI 5 is confìrmed
both for T. longobardicus and T. meridensis. As a quadra-
tojugal is confìrmed to be absent in Tanystropheus (BAI 2,
J43, D35), BA11 (J27), and J39 should be coded not
applicable. Characters BAI, BA5 (J50), BA6, and BAH
(J62) are correctly coded for T. longobardicus and should
be coded thè same for T. meridensis. Dilkes’ coding for
DI 5, overlapping with BA5-6, is accepted. Charac-
ter state 1 (nasals longer than frontals) for BA2 (J61,
DI 8) cannot be evaluated either on thè basis of thè new
material or for T. meridensis. However, PIMUZ T 2484
confirms Benton & Allen’s coding for T. longobardicus ,
while character state 0 (nasals shorter than frontals) for
T. meridensis is rejected. Nasals tapering anteromedially
(J49) or with anterior process at thè midiine (DI 3) can
only be confìrmed on thè basis ot PIMUZ T 2484. Wild
(1974) argued for thè presence of a supratemporal, and
this element is possibly present in MSNM BES SC 265
and in T. meridensis. Both thè presence and interpreta¬
tion of thè supratemporal remain uncertain, however,
and Benton & Allen’s coding for BA13 (D31) is only
provisionally accepted. Jalil’s coding for J14 (D29) for
T. longobardicus (postparietal absent) is accepted and
is conceivably applicable to T. meridensis as well. Any
coding for J3 (postparietal large or small) is consequently
not applicable. Character J63, coded as unknown by Jalil,
is probably better coded 0 (no teeth recurved and later-
ally compressed) as per Rieppel et al. (2003) and Dilkes
(D58-59), both for T. longobardicus and T. meridensis.
Jalil’s coding for skull characters J2, J5, J8, J13, J15-16
(J16, D37), J18, J25 (D8 is described more in detail but
this renders it not applicable to Tanystropheus ), J28, J42,
J68, and J69 is confìrmed, both for T. longobardicus and
T. meridensis. The coding for J17 (large exposure of thè
angular) is correct for T. longobardicus, while it is based
only on Wild’s (1980a) reconstruction for T. meridensis.
However, “large” or “restricted” is subjective. Dilkes’
coding for thè following characters of thè skull is con-
firmed for T. longobardicus : DI, D4-7, D9-10, DI 6- 17,
DI 9-21, D27, D30, D32-34, D36, D46, D49-50, D54-57,
D60-66, D69-72, D75-76, and DI 27. The same character
States as for T. longobardicus are conceivably applicable
to T. meridensis for these characters. Many of Dilkes
characters describe thè dentition and, due to thè pres¬
ence of specialized dentitions in protorosaurs (presence
of tricuspid teeth in Tanystropheus', see also thè dentition
of Langobardisaurus in Renesto & Dalla Vecchia, 2000),
more of these characters should be included in future
analyses of protorosaurs interrelationships. I think that
evidence from MSNM BES SC 1018 and T. meridensis
is consistent with Dilkes’ coding for DI 25 (prefrontals
separate along midiine). Based on thè new specimens of
T. longobardicus and on T. meridensis thè correct charac¬
ter state for D73 is 0. None of thè States described for D74
is applicable to Tanystropheus, in which thè retroarticular
process is formed by thè angular and thè prearticular
without fusion of thè two elements. Dilkes’ coding for
D45 is rejected, because thè state of this character cannot
be assessed in Tanystropheus.
Some characters of thè skull should be coded ditter-
ently in small- and large-sized specimens of Tanystro¬
pheus. Character BA3 is coded 1 (fronto-parietal suture
straight) by Benton & Alien, probably based on thè
large-sized specimen PIMUZ T 2819 but in small-sized
specimens (not clear in T. meridensis) thè suture is inter-
digitating or V-shaped. Character BAI 6, coded 0 (ptery-
goid flange teeth present) for both T. longobardicus and
T. meridensis, should be coded as unknown in thè latter.
The derived state for this character in T. longobardicus
is only applicable to thè large-sized individuate. Dilkes
coded palatine and pterygoid (palatine ramus of) teeth as
absent (D67-68) but this is only true for thè large-sized
specimens (unknown in T. meridensis). By contrast,
TANYSTROPHEUS LONGOBARDICUS'. RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
81
character state for DI 36 (crown of marginai teeth tri-
cuspid) is applicatile only to thè small-sized specimens.
Dilkes’ coding for D25 (parietals fused with loss of
suture) is only applicatile to large-sized Tanystropheus
specimens. The suture is stili partly present in small-
sized specimens or thè two parietals are completely
separated. Dilkes’ coding for D26 (sagittal crest on thè
parietal table present) and D28 (median border of pari-
etal drawn downwards to form ventrolateral flange) is
probably based on thè large PIMUZ T 2819. In thè new
reconstruction of thè skull for thè small-sized specimens
thè parietal table can be described as “constricted with-
out a sagittal crest” and a ventrolateral flange is present
(but see p. 52). However, preservation of thè dorsal sur-
face of thè parietal is very poor in all known small-sized
specimen of Tanystropheus.
The new specimens demonstrate that thè reconstruc¬
tion of thè skull for small-sized specimens of Tanystro¬
pheus is highly speculative in some areas, and that both
Wild’s reconstruction and that presented in this paper
cannot be considered definitive statements on skull mor-
phology. Jalil’s coding for J1 might be correct based on
both reconstructions but thè nature of thè contact nasal-
prefrontal remains hypothetic both for T. longobardicus
and T. meridensis. The coding for 34 and BA9 (J54) is
provisionally accepted for T. longobardicus but thè shape
of thè squamosal and thè nature of its contacts with neigh-
bouring bones have been only tentatively interpreted.
The same characters should be coded as unknown for T.
meridensis. Jalil’s coding for J26 (DI 1-12) (extemal naris
elongate anteroposteriorly and dose to thè midiine) is
probably correct, based only on T. meridensis. Benton &
Allen’s coding for BA8 (posterior process of postorbitai
does not extend beyond back of lower temporal fenestra)
and Dilkes’ coding for D23 (ratio of lengths of anteroven-
tral and posterodorsal processes of postorbitai >1.0) are in
all probability correct for T. longobardicus , while a pos-
torbital was not identified in T. meridensis. A postorbitai
and parietal contact was probably absent (D22, coded
unclearly) and in all probability thè postfrontal entered
into thè margin of thè upper temporal fenestra as stated by
Dilkes (D24). The coding for some characters conceming
thè generai shape of thè skull (J38 and J64, similar to D2)
and thè shape of thè upper temporal fenestra (D3) remains
uncertain.
The coding for some characters of thè skull cannot be
evaluated either on thè basis of thè new material or for
T. meridensis and remains based on Wild’s (1974) recon¬
structions of T. longobardicus. Dilkes’ coding for char¬
acters D38-44, D53, D126, D130, and D142 cannot be
confirmed in thè new specimens because thè bones of thè
dermal palate, thè basicranium, and thè occipital region
are poorly preserved. However, as Wild (1974) thought
that cartilaginous tissue was interposed between thè
paroccipital process and suspensorium, thè nature of their
contact should not be regarded as “strong” (J7), and thè
correct character state for D52 should be 0 (paroccipital
process ends freely). A stape is only tentatively identified
in MSNM BES SC 1018 (Fig. 12) and adds no informa-
tion (see character J6). The prootic is not preserved in any
of thè new specimens and very poorly preserved in thè
PIMUZ material (see characters J70, and D 47-48). Jalil
and Dilkes coded differently thè position of thè occipital
condyle relative to thè craniomandibular joint (J66, ante-
rior to thè craniomandibular joint; D51, even with it) but
as far as can be judged from Wild’s (1974) reconstruction
thè correct coding is that of Dilkes. Although Wild (1974)
stated that a septomaxilla is present in Tanystropheus, I
agree with Dilkes’ coding for D14 as unknown. Evidence
for thè presence of this element seems to be very poor.
The coding for thè following characters of thè axial
skeleton is confirmed or emended. Benton & Allen’s
coding for BAI 7- 19 (J56), BA20 (J40, D82), BA22
(J41), and BA25 (J57, D137) is confirmed for T. longo¬
bardicus. Only thè coding for BA20 and BA22 can be
confirmed for T. meridensis. Character BAI 9 should
be coded as unknown for T. meridensis, as all thè other
characters of thè axial skeleton considered by Benton &
Alien, Jalil, and Dilkes. Trunk intercentra are absent in
T. longobardicus. Consequently, thè coding for BA24 is
rejected, and that for J67 and D80 is confirmed. Char¬
acter BA21 (ovoid spine-table on top of neural spine) is
ambiguous. The shape of thè neural spines of thè dorsal
vertebrae is discussed on pages 62 and 68. Benton &
Allen’s description (BA23=tall and rectangular) is appli¬
cale to thè large-sized specimens of Tanystropheus. For
thè small-sized specimens this character is better captured
by Dilkes (D85), where he codes thè “dorsal neural spine
height” as “low with height < length”. Jalil’s coding for
characters J20, J29 (D83), J36 (D77), J37 is confirmed.
Character J30 should be better described, because only
thè posteriormost cervical ribs have completely separated
tuberculum and capitulum. In thè anterior cervical ribs thè
tuberculum and capitulum are posteriorly confluent and
these ribs are considered to be functionally olocephal-
ous. The character “well-developed transverse processes
of trunk vertebrae” (J31) needs to be re-described. The
transverse processes in thè new specimens from Besano
are short but they might appear shorter than they really
were because of compression (see p. 68). Moreover, while
thè “lumbar” vertebrae supposedly have long pleurapo-
physes, these might altematively be considered as fused
ribs or very long transverse processes (see note on p. 69).
The same is true for thè pleurapophyses of thè proximal
caudals (see Dilkes’ character D89). Jalil’s coding of J19
(no holocephalous dorsal ribs) is rejected. In fact, most of
thè dorsal ribs in Tanystropheus are holocephalous, and
only thè anteriormost three or four are dicocephalous.
This character is better described and correctly coded by
Dilkes (D86). Dilkes’ coding for characters D78-79, D81,
D87, D90, D92, D128, D131-134 is confirmed. “Proximal
caudal neural spine height” (D88) cannot be assessed in
thè new specimens but Dilkes’ coding is applicable to thè
vertebrae of T. conspicuus as described by Wild (1974).
The coding for thè following characters of thè appen-
dicular skeleton is confirmed or emended (all unknown
in T. meridensis). Benton & Allen’s coding for thè
following characters is confirmed: BA26 (J44), BA29
(J47), BA30 (J10, J33 and J51), BA31 (J46), BA33-
35, BA36 (J22, D100), BA37-39, BA40 (J59, DI 15),
BA41 (J35, DI 16), BA42 (J53), BA43 (J58), BA44-45
and BA47 (for BA40 see comments in Rieppel et al.,
2003: 370; for BA41 and BA42 see discussion on pp.
71-72 of this paper). Benton & Allen’s coding of BA32
(metacarpal three equal in length to, or longer than,
fourth) is confirmed, while Jalil’s coding of thè same
character (J55) is rejected. The presence of an entepi-
condylar groove or foramen on thè humerus (BA27)
82
STEFANIA NOSOTTI
was reported by Wild (1974) for T. conspicuus but it
is not confirmed for T. longobardicus (Wild, 1974; see
also thè specimen of T. cf. longobardicus described by
Renesto, 2005), including thè new specimens. Contra
Benton & Alien, both Jalil (J32) and Dilkes (DI 07)
coded an entepicondylar foramen absent. Character
state 0 for BA28 (radius length relative to thè humerus)
is questionable. In thè well preserved specimens
described here thè radius is 66-70% thè length of thè
humerus. The desumption of thè shape of metatarsal V
(BA46, J12, DI 22) is a character stili causing confu-
sion (Rieppel et al., 2003: 370), and clearer descrip-
tions are needed. Jalil’s coding is confirmed for J9
(D93), Jll (D121), J23, J24 (D114), J45, J52, and J71
(D 1 38). An ectepicondylar foramen was never reported
for Tanystropheus. Both Jalil (J21) and Dilkes (DI 08)
coded it as absent. I reject Jalil ’s coding for J48 (ilium
with reduced contribution in thè acetabulum), and
Dilkes’ coding for DI 05 (relative contribution of pubic
elements to acetabulum approximately equal) because
thè new specimens demonstrate that thè element pri-
marily contributing to thè acetabulum is thè ilium (Fig.
25). A concave-convex astragalo-calcaneum articula-
tion is definitely absent in Tanystropheus (see discus-
sion on p. 71): Jalil’s coding for J34 is rejected, and
Dilkes’ coding for DI 13 confirmed. Dilkes’ coding is
confirmed for D94, D97, D99, D101-102, D104, D106,
D109, DI 12, DI 19-120, D123-124, D135 and D144.
Dilkes’ coding for characters describing thè shape of
thè interclavicle (D96 and D98) is based on Wild’s
interpretation of a single, fragmentary element (Wild,
1974: fig. 64). An interclavicle is only tentatively iden-
tifìed in MSNM BES SC 1018 (Fig. 20). The presence
of a processus lateralis on thè pubis (DI 03) established
by Wild (1974) is not confirmed in thè new specimens
(p. 34). Presence and position of a centrale (D 1 1 7- 1 1 8)
is correctly coded by Dilkes as unknown. The presence
of a cartilaginous centrale in Tanystropheus remains
highly conjectural (see discussion on p. 72). However,
an ossified centrale contacting thè tibia is present in thè
related taxa Langobardisaurus and Macrocnemus.
Finally, coding for BA48 (J60) (postcloacal bones)
cannot be confirmed in thè new specimens but thè pres¬
ence of postcloacal bones in Tanystropheus was estab¬
lished by Wild (1974) in thè PIMUZ material.
At present, there is a generai consensus on thè archo-
sauromorph, particularly Archosauria, affinities of Tanys¬
tropheus. The inclusion of Protorosauria (thè priority of
Protorosauria Huxley, 1871 over Prolacertiformes was
established by Chatterjee, 1986) within thè Archosauro-
morpha was formerly put forward by Gow (1975), and
was Consolidated in thè 1980ies through thè work of
several palaeontologists, exhaustively summarized by
Benton (1985) and Evans (1988). The strongest argu-
ments against thè inclusion of thè Protorosauria inside thè
Archosauromorpha remain those of Wild (1974; 1980 a).
In a preliminary paper (Nosotti, 1999), I maintained that
thè tiny elements observed in thè elbow and thè knee
joints in MSNM BES SC 265 should be interpreted
as secondary epiphyseal centers of ossifìcation. This
character was recognized by Benton (1985) and Evans
(1988) as a lepidosaurian synapomorphy but was also
described in pterodactyloid pterosaurs (Bennett, 1993),
and discussed in thè protorosaur Macrocnemus (Premru,
1991; Rieppel, 1989). Through a closer examination of
these elements and taken together with thè nature of thè
articular ends of thè long bones in thè new Besano speci¬
mens, I concluded that my earlier interpretation should
probably be rejected.
Dilkes’ ( 1 998) cladistic analysis of protorosaurs inter-
relationships recently raised thè possibility that Proto¬
rosauria as conventionally conceived are paraphyletic.
This result was confirmed by thè re-analysis of Rieppel
et al. (2003). This latter paper should be consulted for an
update on thè more recent views on thè relationships of
Tanystropheus with other protorosaurian taxa. The sister
group relationships of Tanystropheus and Tanytrachelos
(Olsen, 1979), thè two being thè representatives of thè
Family Tanystropheidae, is, however, unanimously con¬
firmed in all cladistic analyses that include both these taxa
(Benton, 1985; Evans, 1988; Jalil, 1994; Benton & Alien,
1997; Peters, 2000a; Rieppel et al., 2003). Tanytrachelos
ahynis, from thè Upper Triassic (Camian) of thè USA
was interpreted by Olsen (1979) as an aquatic animai. At
present, Tanytrachelos remains poorly described. More
information on thè anatomy of Tanytrachelos would
provide thè opportunity for interesting comparisons with
Tanystropheus and perhaps offer new insights into thè
supposed aquatic radiation of thè (terrestrial) protorosaurs
represented by thè Tanystropheidae.
CONCLUSIONS
1) New material from thè Middle Triassic of Besano
adds considerable detail to thè knowledge of thè anatomy
of Tanystropheus longobardicus. Re-interpretations ofthe
anatomy presented in this paper are only applicable to
small-sized representatives of T. longobardicus and also
to T. meridensis, that cannot be distinguished from thè
smallest specimens of T. longobardicus and is regarded
here as conspecific. Three larger-sized Tanystropheus
specimens are comprised in thè new material and their
morphology is described and discussed.
2) A new reconstruction ofthe skull is presented, devel-
oped from a clay three-dimensional model. Major novel-
ties in thè interpretation of thè pre-orbital region concem
thè nature of thè contacts between thè different elements
and thè shape of thè prefrontals. The shape, location and
contacts of thè nasals are reconstructed by comparison
with Tanystropheus material in thè PIMUZ Collections
and remain partly speculative. The contacts between
thè nasals, prefrontals and frontals are only tentatively
reconstructed. The fronto-parietal piate is re-interpreted:
thè lateral flanges of thè frontal are horizontally rather
than vertically oriented and thè features distinguishing
thè dorsal and ventral surfaces of thè fronto-parietal piate
are described. Major difficulties were encountered in thè
reconstruction of thè temporal region: thè postorbitai and
thè squamosal are re-described and a new tentative inter¬
pretation of their contacts with thè neighbouring bones is
given. The presence of a sclerotic ring in thè orbit is defi¬
nitely confirmed. The shape and contacts of thè elements
of thè lower jaw are partly re-interpreted.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRET ATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
83
Fig. 67 - Tanystropheus longobardicus . Watercolor: Fabio Fogliazza.
tì uZ ’%
84
STEFANIA NOSOTTI
3) A re-description of thè axial skeleton is given,
mainly based on thè fully articulated MSNM BES SC
265. Differences in vertebral morphology between small-
and large-sized specimens of Tanystropheus are assessed
through comparison with three-dimensionally preserved
vertebrae referred to Tanystropheus conspicuus and
Tanystropheus cf. longobardicus.
4) The superbly preserved limbs of MSNM BES SC
1018 provide for thè first time unequivocal information
on thè elements of thè manus and pes and their articular
relationships. A re-description of thè tarsus based on
thè new specimens is given. The presence of sesamoid
bones in thè elbow and knee joints is reported for thè
first time.
5) Discussion on thè morphology of thè hindlimb
highlights thè difficulties in thè interpretation of ter-
restrial locomotion in Tanystropheus. It is concluded
that, although thè morphology of thè hindlimb is not
strikingly specialized for locomotion in water, it is
more consistent with an aquatic mode of fife for Tanys¬
tropheus.
6) Based on thè overall skeletal anatomy, Tanystroph¬
eus is regarded as an aquatic protorosaur with dose ter-
restrial ancestors, living in shallow waters. At present, thè
more consistent model for aquatic locomotion in Tanys¬
tropheus seems to be that suggested by Tschanz (1985;
1986), who envisaged Tanystropheus as an axial-subun-
dulational swimmer relying on lateral undulations of thè
trunk and tail for aquatic propulsion, perhaps enhanced
by paddling with thè hindlimbs. Major difficulties are
encountered in thè assessment of thè method of capture
and processing thè prey adopted by Tanystropheus. The
cheek dentition of extant pinnipeds is proposed as a new
model for thè understanding of thè functional significance
of thè tricuspid dentition in Tanystropheus.
7) The notion that thè extremely elongated neck of
Tanystropheus was inadaptive is rejected, based also on
comparison with plesiosauroid plesiosaurs.
8) Description and coding of thè phylogenetically
informative characters used in thè most recent cladistic
analyses are discussed in thè light of thè new information
provided by thè new material.
Acknowledgements
I am deeply grateful to thè volunteers of thè Paleon-
tological Group of Besano who excavated in thè Besano
quarries for thè last three decades. They provided me with
thè superbly preserved material which first gave me thè
opportunity to undertake this exciting enterprise.
I warmly thank: thè Soprintendenza per i Beni
Archeologici della Lombardia for assistance in obtaining
permission for excavations at thè Sasso Caldo site near
Besano; thè Regione Lombardia, Assessorato alle Cul¬
ture, Identità, Autonomie della Lombardia for financial
support for thè preparation of thè specimens and research;
Giovanni Tafuni (Ospedale Maggiore di Milano) for
thè radiographs; Giuseppina Damiano (MSNM), Fabio
Fogliazza (MSNM) and Sergio Rampinelli for thè skil-
ful preparation of thè specimens; Roberto Appiani, Mas¬
simo Demma, Nicholas C. Fraser, Heinz Lanz (PIMUZ),
Rosi Roth (PIMUZ) and Luciano Spezia (MSNM) for
thè photographs; Massimo Demma and Fabio Fogliazza
(MSNM) for thè line drawings, watercolors and pencils;
Massimo Demma, Simone Maganuco and Michela Mura
for thè editing of illustrations; Kathleen Histon for thè
translation of thè dedication; Giacomo Bracchi (MSNM
and Museo Civico di Storia Naturale di Piacenza) for
careful reading of thè proofs.
I extend my warmest thanks to: Hugo Bucher and
Heinz Furrer (PIMUZ) for allowing me access to thè
specimens in their Collections and Heinz Lanz (PIMUZ)
for his generous assistance and hospitality during my visit
in Zurich; Rudolf Stockar (Museo Cantonale di Storia
Naturale, Lugano) for allowing me access to specimen
MCSN 4451 in his care.
I am deeply indebted to thè reviewers Nicholas
C. Fraser (Virginia Museum of Naturai History, Martin-
sville), Simone Maganuco (MSNM and Dipartimento di
Scienze della Terra, Università degli Studi di Firenze)
and Olivier Rieppel (Field Museum of Naturai History,
Chicago) for comments, criticism and discussion, which
greatly improved thè originai manuscript. N. C. Fraser
and O. Rieppel also edited thè English text, greatly
improving its style and thè clarity of thè contents. I thank
S. Maganuco also for thè generous and Constant support,
which helped me to overcome thè numerous difficulties
encountered in completing my job.
Silvio Renesto (Università degli Studi dell’Insubria,
Varese) provided helpffil comments and criticism on a
very early draft of this paper.
My interpretation of thè temporal region of thè skull
was greatly improved through discussion with N. C. Fraser
and S. Maganuco. I profìted also ffom discussion with
Cristiano Dal Sasso (MSNM), who offered valuable sug-
gestions and hints for further investigations. Likewise I am
indebted to Richard Forrest for discussion on thè mode of
life of Tanystropheus, for sharing information on his origi¬
nai work, and giving access to unpublished data and litera-
ture on plesiosaurs. Giovanni Pasini (Museo dei Fossili di
Besano) offered helpful suggestions and was always gener-
ously available for discussion. Over thè years I have greatly
benefited ffom discussions with Rupert Wild (Staatliches
Museum furNaturkunde, Stuttgart). His outstanding mono-
graph on Tanystropheus was an important reference for my
job. Giuseppe Muscio (MFSN) shared with me information
on his originai work. The friend Marco Auditore provided
much needed access to literature.
I thank for help and discussion David Martill (Uni¬
versity of Portsmouth) and Lars Schmidt (Department
of Geology, University of California), as well as thè fol-
lowing colleagues from MSNM: Giorgio Bardelli, Valter
Fogato, Paola Livi, Maurizio Pavesi, Michela Podestà,
Fabrizio Rigato, Stefano Scali.
My hearthly thank to Michela Mura (Graphic design,
MSNM) for her friendly assistence and skilful editing of
this monograph.
I acknowledge financial support for thè publication of
this monograph from Cinehollywood s.r.l.
Linguistic mistakes, errors of fact or interpretation,
along with any inadvertent omissions are my own.
TANYSTROPHEUS LONGOBARDICUS: RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
85
REFERENCES
Arthaber G. v., 1921 - Studien iiber Flugsaurier auf
Grund der Bearbeitung des Wiener Exemplares von
Dorygnathus Banthensis Theod. sp. Denkschriften
der Akademie der Wissenschaften in Wien, Mathema-
tisch-Naturwissenschaftliche K lasse, Wien, 97 (1919):
391-464.
Arthaber G. v., 1922 - Uber Entwicklung, Ausbildung
und Absterben der Flugsaurier. Palàontologische Zeit-
schrift , Berlin, 4: 1-47.
Avanzini M. & Renesto S., 2002 - Areview of Rhyncho-
sauroides tirolicus Abel, 1926 ichnospecies (Middle
Triassici Anisian-Ladinian) and some inferences on
Rhynchosauroides trackmaker. Rivista Italiana di
Paleontologia e Stratigrafia, Milano, 108 (1): 51-
66.
Bassani F., 1886 - Sui fossili e sull’età degli schisti bitu¬
minosi triasici di Besano in Lombardia. Atti Società
Italiana di Scienze Naturali, Milano, 29: 15-72.
Bennett S. C., 1993 - The ontogeny of Pteranodon and
other pterosaurs. Paleobiology, Chicago, 19 (1): 92-
106.
Benton M. J., 1985 - Classification and phylogeny of thè
diapsid reptiles. Zoologica! Journal of thè Linnean
Society, London, 84: 97-164.
Benton M. J. & Alien J. L., 1997 - Boreopricea from thè
Lower Triassic of Russia, and thè relationships of thè
prolacertiform reptiles. Palaeontology, London, 40:
931-953.
Braun J. & Reif W.-E., 1982 - A new terminology of
aquatic propulsion in vertebrates. Neues Jahrbuch fiir
Geologie und Palàontologie-Abhandlungen, Stuttgart,
164(1-2): 162-167.
Braun J. & Reif W.-E., 1985 - A survey of aquatic loco-
motion in fìshes and tetrapods. Neues Jahrbuch fiir
Geologie und Palàontologie-Abhandlungen, Stuttgart,
169(3): 307-332.
Brinkman D., 1980 - Structural correlates of tarsal and
metatarsal fimctioning in Iguana (Lacertilia; Iguani-
dae) and other lizards. Canadian Journal of Zoology,
Ottawa, 58: 277-289.
Brinkman D., 1981 - The origin of thè crocodiloid tarsi
and thè interrelationships of thecodontian archosaurs.
Breviora Museum of Comparative Zoology, Cam¬
bridge, Massachusetts, 464: 1-23.
Broili F., 1915 - Beobachtungen an Tanystropheus cons-
picuus H. v. Meyer. Neues Jahrbuch fiir Mineralogie,
Geologie und Palàontologie, Stuttgart, II: 51-62.
Carroll R. L. & Gaskill P., 1985 - The nothosaur Pachyp-
leurosaurus and thè origin of plesiosaurs. Philosophi-
cal Transactions of thè Royal Society, London, B 309
(1139): 343-393.
Chapple D. G. & Swain R., 2002 - Distribution of energy
reserves in a viviparous shink: does tail autotomy
involve thè loss of lipid Stores? Austro/ Ecology, 27
(5): 565-572.
Chatterjee S., 1986 - Malerisaurus langstoni, a new diap¬
sid reptile from thè Triassic of Texas. Journal of Verte¬
brate Paleontology, Northbrook, 6 (4): 297-312.
Cox B., 1975 - The longest-necked lizard? Nature,
London, 254: 654-655.
Dalla Vecchia F. M., 2000 - Tanystropheus (Archosauro-
morpha, Prolacertiformes) remains from thè Triassic of
thè Northern Friuli (NE Italy). Rivista Italiana di Pale-
ontologia e Stratigrafia, Milano, 106(2): 135-140.
Dalla Vecchia F. M., 2006 - Resti di Tanystropheus, Sau-
rotterigi e “Rauisuchi” (Reptilia) nel Triassico medio
della Val Aupa (Moggio Udinese, Udine). Gortania
- Atti del Museo Friulano di Storia Naturale, Udine,
27 (2005): 25-48.
Dalla Vecchia F. M. & Avanzini M., 2002 - New scat-
tered remains of Triassic reptiles from northeastem
Italy. Bollettino della Società Paleontologica Italiana,
Modena, 41 (2/3): 215-235.
Demes B. & Krause D. W., 2005 - Suction feeding in a
Triassic protorosaur? Science, Washington, 308: 1112-
1113.
Dilkes D. W., 1998 - The Early Triassic rhynchosaur
Mesosuchus browni and thè interrelationships of
basai archosauromorph reptiles. Philosophical
Transactions of thè Royal Society, London, B 353
(1368): 501-541.
Edmund A. G., 1969 - Dentition. In: The Biology of thè
Reptilia. Gans C. (ed.). Academic Press, London &
New York, 1: 117-200.
Ellenberger P., 1977 - Quelques précisions sur l’anato-
mie et la place systématique trés spéciale de Cose-
saurus aviceps. (Ladinien supérieur de Montrall,
Catalogne). Cuadernos de Geologia Ibèrica, Madrid,
4: 169-188.
Evans S., 1988 - The early history and relationships of
thè Diapsida. In: The phylogeny and classification of
thè tetrapods. Benton M. J. (ed.). Clarendon Press,
Oxford, 1: 221-260.
Femàndez M. & Gasparini Z., 2000 - Salt glands in a
Tithonian metriorhynchid crocodyliform and their
physiological signifìcance. Lethaia, Oslo, 33: 269-
276.
Ford T., 2002 - A new interpretation of thè skull of Tanys¬
tropheus. Journal of Vertebrate Paleontology, North¬
brook, 22 (3, supplement): 53A.
Fraser N. C., Nosotti S. & Rieppel O., 2004 - A re-evalu-
ation of two species of Tanystropheus (Diapsida, Pro-
torosauria), from Monte San Giorgio, southern Alps.
Journal of Vertebrate Paleontology, Hanover, 24 (3,
supplement): 60A.
Fraser N. C. & Rieppel O., 2006 - A new protorosaur
(Diapsida) from thè Upper Buntsandstein of thè Black
Forest, Germany. Journal of Vertebrate Paleontology,
Northbrook, 26 (4): 866-871.
Furrer H., 1995 - The Kalkschieferzone (Upper Meride
Limestone; Ladinian) near Meride (Canton Ticino,
Southern Switzerland) and thè evolution of a Middle
Triassic intraplatform basin. Eclogae geologicae Hel-
vetiae, Basel, 88: 827-852.
Gauthier J., Kluge A. G. & Rowe T., 1988 - Amniote
phylogeny and thè importance of fossils. Cladistics,
London, 4: 105-209.
Gould S. J., 1974 - The origin and function of “bizarre”
structures: antler size and skull size in thè “Irish Elk”,
Megaloceros giganteus. Evolution, 28: 191-220.
Gow C. E., 1975 - The morphology and relationships
of Youngina capensis Broom and Pro/acerta broomi
Parrington. Paleontologia Africana, Johannesburg,
18: 89-131.
86
STEFANIA NOSOTTI
Hopson J. A., 1979 - Paleoneurology. In: Biology of thè
Reptilia. Gans C. (ed.). Academic Press, London, 9A:
39-146.
Huene F. v., 1907-1908 - Die Dinosauriers der Euro-
pàischen Triasformation, mit Beriicksichtigung der
aussereuropaischen Vorkommnisse. Geologische und
Pa/aeontologische Abhandlungen, Supplement-Band
1 . Gustav Fischer, Jena.
Huxley T. H., 1871 - A Manual of thè Anatomy of Verte¬
brate Animals. J. & A. Churchill, London.
I.C.Z.N., 1981 - Opinion 1186. Tanystropheus H. v.
Meyer, [1852] (Reptilia) conserved. Bulletin of
Zoological Nomenclature, London, 38 (3): 1 88-
190.
Jalil N.-E., 1997 - A new prolacertifomi diapsid from thè
Triassic of North Africa and thè interrelationships of
thè Prolacertiformes. Journal of Vertebrate Paleontol-
ogy, Northbrook, 17 (3): 506-525.
Jurcsàk T., 1975 - Tanystropheus biharicus n. sp. (Rep¬
tilia, Squamata) o nouà specie pentru fauna Triasicà a
Romàniei. Nymphaea, Oradea, 3: 45-52.
Jurcsàk T., 1 976 - Noi descoperiri de reptile fosile in Tria-
sicul de la Ale§d. Nymphaea, Oradea, 4: 67-105.
Jurcsàk T., 1978 - Rezultate noi in studiul saurienilor
fosili de la Ale§d (Bihor, Romania). Nymphaea,
Oradea, 6: 15-60.
Jurcsàk T., 1982 - Occurrences nouvelles des Sauriens
mèsozoiques de Roumania. Vertebrata Fiungarica,
Budapest, 21: 175-184.
Kuhn O., 1934 - Sauropterygia. In: Fossilium catalogus.
I, Ammalia. Quenstedt W. (ed.). W Junk., Berlin, Pars
69.
Kuhn O., 1963 - Sauria. In: Fossilium catalogus. I, Ani-
malia. Westphal F. (ed.). W Junk., Berlin, Pars 104
(Supplementum I).
Kuhn-Schnyder E., 1960 - Hand und Fuss von Tanystro¬
pheus longobardicus (Bassani). Eclogae geologicae
Helvetiae, Basel, 52 (1959) (2): 921-941.
Kuhn-Schnyder E., 1967 - Das Problem der Euryapsida.
Mitteilungen aus dem Palàontologischen Institut der
Universitàt Zurich, Zurich, 49: 335-348.
Kummer B., 1975 - Biomechanik fossiler und rezenter
Wirbeltiere. Natur und Museum, Frankfurt am Main,
105: 156-167.
LaBarbera M. & Rieppel O., 2005 - Suction feeding in a
Triassic protorosaur? Science, Washington, 308: 111 2-
1113.
Li Chun, 2003 - First record of Tanystropheid Reptile
(Order Protorosauria) from thè Middle Triassic of
China. Acta Geologica Sinica (English Edition), Bei¬
jing, Il (4): 419-423.
Li Chun, Rieppel O. & LaBarbera M. C., 2004 - A triassic
aquatic protorosaur with an extremely long neck. Sci¬
ence, Washington, 305 (5692): 1931.
Long J. H. Jr., Schumacher J., Livingston N. & Kemp M.,
2006 - Four flippers or two? Tetrapodal swimming
with an aquatic robot. Bioinspiration & Biomimetics,
Bristol, 1 : 20-29.
Massare J. A., 1 988 - Swimming capabilities of Mesozoic
marine reptiles: implications for method of predation.
Paleobiology, Chicago, 14(2): 187-205.
McHenry C. R., Cook A. G. & Wroe S., 2005 - Bottom-
feeding plesiosaurs. Science, Washington, 310 (5745):
75.
Meyer H. v., 1 847- 1 855 - Die Saurier des Muschelkalkes:
mit Rucksicht auf die Saurier aus Buntem Sandstein
und Keuper. In: Zur Fauna der Vorwelt, zweite Abthei-
lung. Heinrich Keller, Frankfurt a.m.
Miyazaki N., 2002a - Teeth. In: Encyclopedia of
Marine Mammals. Perrin W. F., Wiirsig B. & The-
wissen J. G. M. (eds.). Academic Press, San Diego:
1227-1232.
Miyazaki N., 2002b - Ringed, Caspian, and Baikal Seals.
In: Encyclopedia of Marine Mammals. Perrin W. F.,
Wiirsig B. & Thewissen J. G. M. (eds.). Academic
Press, San Diego: 1033-1037.
Modesto S. P. & Sues H-D., 2004 - The skull of thè Early
Triassic archosauromorph reptile Pro/acerta broomi
and its phylogenetic significale. Zoological Journal
of thè Linnean Society, London, 140: 335-351.
Motani R., 1997 - Temporal and spatial distribution
of tooth implantations in ichthyosaurs. In: Ancient
Marine Reptiles. Callaway J. M. & Nicholls E. L.
(eds.). Academic Press, 81-103.
Munster G. zu, 1834 - Vorlàufìge Nachricht iiber einige
neue Reptilien im Muschelkalke von Baiem. Neues
Jahrbuch fur Mineralogie, Geognosie, Geologie und
Petrefaktenkunde, Stuttgart: 521-526.
Muscio G., 1 997 - Preliminary note on a specimen of Pro¬
lacertiformes (Reptilia) from thè Norian (Late Trias¬
sic) of Preone (Udine, North-Eastem Italy). Gortania
- Atti del Museo Friulano di Storia Naturale, Udine,
18(1996): 33-40.
Noè L., 2006 - The role of thè plesiosaurian long neck.
A new model. Journal of Vertebrate Paleontology,
Northbrook, 26 (3, supplementi 105A.
Nopcsa F. v., 1923 - Neubeschreibung des Trias-Pte-
rosauriers Tribelesodon. Palàontologische Zeitschrift,
Berlin, 5: 161-181.
Nosotti S., 1999 - New fìndings of Tanystropheus longo¬
bardicus (Reptilia, Prolacertiformes) in thè Middle
Triassic of Besano (Lombardy, Northern Italy). In:
Third International Symposium on lithographic lime-
stones. Renesto S. (ed.). Rivista del Museo civico di
Scienze Naturali “E. Caffi”, Bergamo, 20 (supple¬
mento): 1 17-120.
Oelrich T. M., 1956 - The Anatomy of thè Head of Cteno-
saura pectinata (Iguanidae). Miscellaneous Publica-
tions, Museum of Zoology, University of Michigan,
94: 9-122.
O’Keefe F. R. & Robin F., 2001 - Acladistic analysis and
taxonomic revision of thè Plesiosauria (Reptilia: Sau¬
ropterygia). Acta Zoologica Fennica, 213:1 -63.
Olsen P. E., 1979 - A new aquatic Eosuchian from thè
Newark Supergroup (Late Triassic-Early Jurassic) of
North Carolina and Virginia. Postilla, New Haven,
176: 1-14.
Ortlam D., 1967 - Fossile Bòden als Leithorizonte fur die
Gliederung des Hòheren Buntsandsteins im nòrdlichen
Schwarzwald und siidlichen Odenwald. Geologisches
Jahrbuch, Hannover, 84: 485-590.
Peters D., 2000a - A reexamination of four prolacer-
tiforms with implications for pterosaur phylogen-
esis. Rivista Italiana di Paleontologia e Stratigrafia,
Milano, 106 (3): 293-336.
Peters D., 2000b - Description and interpretation of inter-
phalangeal lines in tetrapods. Ichnos, Chur & New
York, 7 (1): 11-41.
TANYSTROPHEUS LONGOBARDICUS-, RE-INTERPRETATIONS OF THE ANATOMY BASED ON NEW SPECIMENS FROM BESANO
87
Peters D., 2005 - Suction feeding in a Triassic protoro-
saur? Science , Washington, 308: 1112.
Peyer B., 1 93 1 - Die Triasfauna der Tessiner Kalkalpen II.
Tanystropheus longobardicus Bass. sp. Abhandlungen
der Schweizerischen Palaontologìschen Gesellschaft ,
Basel, 50: 9-110.
Premru E., 1991 - Beschreibung eines neuen Fundes von
Macrocnemus bassanii Nopcsa (Reptilia, Squamata,
Prolacertiformes) aus der Grenzbitumenzone (Anis/
Ladin) von Besano, Italien. Diplomarbeit an der
Philosophischen Fakuìtdt II der Universitdt Zurich,
Ziirich: 1-57.
Quenstedt W., 1963 -Clavis bibliographica. In: Fossilium
catalogus. I, Ammalia. Westphal F. (ed.). W. Junk.,
Berlin, Pars 102.
Renesto S., 1994 - A new prolacertiform reptile from
thè Late Triassic of Northern Italy. Rivista Italiana
di Paleontologia e Stratigrafia, Milano, 100 (2): 285-
306.
Renesto S., 2005 - A new specimen of Tanystropheus
(Reptilia Protorosauria) from thè middle Triassic of
Switzerland and thè ecology of thè genus. Rivista
Italiana di Paleontologia e Stratigrafia, Milano, 1 1 1
(3): 377-394.
Renesto S. & Avanzini M., 2002 - Skin remains in a juve-
nile Macrocnemus bassanii Nopcsa (Reptilia, Prolac¬
ertiformes) from thè Middle Triassic of Northern Italy.
Neues Jahrbuch fiir Geologie und Palàontologie.
Abhandlungen, Stuttgart, 224 (1): 31-48.
Renesto S. & Dalla Vecchia F. M., 2000 - The unusual
dentition and feeding habits of thè prolacertiform rep¬
tile Langobardisaurus (Late Triassic, Northern Italy).
Journal of Vertebrate Paleontologv, Northbrook, 20
(3): 622-627.
Renesto S., Dalla Vecchia F. M. & Peters D., 2002
- Morphological evidence for Bipedalism in thè Late
Triassic Prolacertiform Reptile Langobardisaurus.
Senckenbergiana lethaea, Frankfurt am Main, 82 (1):
95-106.
Rewcastle S. C., 1980 - Form and function in lacertil-
ian knee and mesotarsal joints; a contribution to thè
analysis of sprawling locomotion. Journal ofZoology,
London, 191: 147-170.
Rieber H., 1973 - Cephalopoden aus der Grenzbitumen¬
zone (Mittlere Trias) des Monte San Giorgio (Kanton
Tessin, Schweiz). Schweizerischen Pai dnto/ogischen
Abhandlungen , Basel, 93: 1-96.
Rieppel O., 1989 - The hind limb of Macrocnemus bas¬
sanii (Nopcsa) (Reptilia, Diapsida): development and
functional anatomy. Journal of Vertebrate Pa/eontol-
ogy, Northbrook, 9 (4): 373-387.
Rieppel O., 2000 - Middle Triassic marine vertebrates
from thè northern Gondwanan shelf. Zentralblatt fiir
Geologie und Palàontologie, Stuttgart, I (9-10): 1269-
1284.
Rieppel O., 2001 - A new species of Tanystropheus
(Reptilia: Protorosauria) from thè Middle Triassic of
Makhtesh Ramon. Israel. Neues Jahrbuch fiir Geolo¬
gie und Palàontologie. Abhandlungen, Stuttgart, 221
(2): 271-287.
Rieppel O. & Gronowski, 1981 - The loss of thè lower
temporal arcade in thè diapsid reptiles. Zoologica!
Journal of thè Linnean Society, London, 72: 203-
217.
Rieppel O., Fraser N. C. & Nosotti S., 2003 - The mono-
phyly of Protorosauria (Reptilia, Archosauromorpha):
a preliminary analysis. Atti della Società italiana di
Scienze naturali e Museo civico di Storia naturale in
Milano, Milano, 144 (II): 359-382.
Robinson P. L., 1975 - The functions of thè hooked fifth
metatarsal in lepidosaurian reptiles. Colloque interna-
tional C.N.R.S. , Paris, 218: 461-483.
Ròhl H. J., Schmid-Ròhl A., Furrer H., Frimmel A., Osch-
mann W. & Schwark L., 2001 - Microfacies, geoche-
mistry and palaeoecology of thè Middle Triassic
Grenzbitumenzone from Monte San Giorgio (Canton
Ticino, Switzerland). Geologia Insubrica, Lomazzo,
6(1): 1-13.
Romer A. S., 1956 - Osteology of thè Reptiles. The Uni¬
versity of Chicago Press, Chicago.
Sanz J. L. & López-Martinez N., 1984 - The prolacertid
lepidosaurian Cosesaurus aviceps Ellenberger & Vil¬
lana, a claimed “protoavian” from thè Middle Triassic
of Spain. Geobios, Lyon, 17: 747-753.
Schaeffer B., 1941 - The morphological and functional
evolution of thè tarsus in amphibians and reptiles.
Bulletin of thè American Museum of Naturai History,
New York, 78 (6): 395-472.
Scheffer V. B., 1969 - Seals, Sea Lions, and Walruses.
Stanford University Press, Stanford.
Senter P., 2007 - Necks for sex: sexual selection as an
explanation for sauropod dinosaur neck elongation.
Journal ofZoology, London, 271 (1): 45-53.
Simmons R. E. & Scheepers L., 1996 - Winning by a
neck: sexual selection in thè evolution of giraffe.
The American Naturalist, Chicago, 148 (5): 77 1 -
786.
Smith A. S., 2003 - Cladistic analysis of thè Plesiosauria
(Reptilia: Sauropterygia). M. Se. unpublished Thesis,
University of Bristol, Department of Earth Sciences,
Faculty of Sciences.
Sulej T., 2004 - Krasiejow. The remarkable discovery of
Triassc pre-dinosaurs. Przygod Studio, Opole.
Taylor M. A., 1989 - Neck and neck. Nature, London,
341 (6244): 688-689.
Tschanz K., 1985 - Tanystropheus - An unusual reptilian
construction. In: Konstruktionsprinzipien lebender
und ausgestorbener Reptilien. Konzepte SFB 230.
Stuttgart, (4): 169-178.
Tschanz K., 1986 - Funktionelle Anatomie der Halswir-
belsàule von Tanistropheus longobardicus (Bassani)
aus der Trias (Anis/Ladin) des Monte San Giorgio
(Tessin) auf der Basis vergleichend morphologischer
Untersuchungen an der Halsmuskulatur rezenter
Echsen. Ph. D. Thesis Universitdt Zurich, AD AG
Administration & Druck AG., Zurich.
Tschanz K., 1988 - Allometry and heterochronic proc-
esses in thè neck-growth of thè Triassic prolacertiform
reptiles Tanystropheus and Macrocnemus. Palaeontol-
ogy, London, 31: 153-179.
Underwood G., 1970 - The Eye. In: The Biology of thè
Reptilia. Gans C. & Parson T. S. (eds.). Academic
Press, London & New York, 2: 1-97.
Vickers-Rich P., Rich T., Rieppel O., Thulborn R. A. &
McClure H. A., 1999 - A Middle Triassic vertebrate
fauna from thè Jilh Formation, Saudi Arabia. Neues
Jahrbuch fiir Geologie und Palàontologie Abhandlun¬
gen, Stuttgart, 213 (2): 201-232.
88
STEFANIA NOSOTTI
Wild R., 1974 - Die Triasfauna der Tessiner Kalkalpen
XXIII. Tanystropheus longobardicus (Bassani) (neue
Ergebnisse). Schweizerische Palàontologische Abhan-
dlungen, Basel, 95 (1973): 4-162.
Wild R., 1975 - Tanystropheus H. v. Meyer, 1855 (Rep-
tilia): request for conservation under thè plenary
powers. Z.N.(S.) 2084. Bulletin ofZooIogical Nomen¬
clature, London, 32 (2): 124-126.
Wild R., 1976 - Tanystropheus H. v. Meyer, [1852]
(Reptilia): revised request for conservation under
thè plenary powers. Z.N.(S.) 2084. Bulletin of
Zoologica! Nomenclature, London, 33 (2): 124-
126.
Wild R., 1980a - Die Triasfauna der Tessiner Kalkal¬
pen XXIV. Neue Lunde von Tanystropheus (Rep¬
tilia, Squamata). Schweizerische Palàontologische
Abhandlungen, Basel, 102: 4-43.
Wild R., 1980b - Tanystropheus (Reptilia: Squamata) and
its importance for stratigraphy. Mémoires de la Société
Géologique de France, n.s., Paris, 139: 201-206.
Wild R., 1987 - An example of biological reasons for
extinction: Tanystropheus (Reptilia, Squamata).
Mémoires de la Société Géologique de France, n. s.,
Paris, 150: 37-44.
Wild R. & Oosterink H., 1984 - Tanystropheus (Reptilia,
Squamata) aus dem Unteren Muschelkalk von Win-
terswijk, Holland. Grondboor en Hamer, Oldenzaal,
5: 142-148.
Zaher H. & Rieppel O., 1999 - Tooth implantation and
replacement in squamates, with special reference to
mosasaur lizard and snakes. American Museum Novi-
tates, New York, 3271: 1-19.
Zamik B., 1925 - K ethologiji plesiosaurija, sa prinosima
mehanici kraljeznice u recentnih sauropsida. Zagreb.
Stefania Nosotti - Museo Civico di Storia Naturale di Milano, Sezione di Paleontologia, Corso Venezia 55, 20121 Milano, Italy.
e-mail: stefanianosotti@yahoo.it
Tanystropheus longobardicus (Reptilia, Protorosauria): re-interpretations ot thè anatomy based on new specimens troni thè Middle
Triassic of Besano (Lombardy, northem Italy)
Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano
Volume XXXV - Fascicolo III
.
NOSOTTI S. -
Tanystropheus ìongobardicus
(Reptilia, Protorosauria):
re-interpretations of thè anatomy
based on new specimens from
thè Middle Triassic of Besano
(Lombardy, northem Italy)
*
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I
Memorie Soc. It. Sci Nat. e
Museo Civ. St. Nat. Milano
Volume XXXV - Fascicolo III
Piate I - MSNM BES SC 265
Scale bar 50 mm.
Preparation: Sergio Rampinelli.
Photo: Luciano Spezia.
NOSOTTI S. -
Tanystropheus longobardicus
(Reptilia, Protorosauria):
re-interpretations of thè anatomy
based on new specimens from
thè Middle Triassic of Besano
(Lombardy, northem Italy)
Memorie Soc. It. Sci Nat. e
Museo Civ. St. Nat. Milano
Volume XXXV - Fascicolo III
Piate II - MSNM BES SC 265
Drawing: Fabio Fogliazza.
►SOTTI S.
Tanystropheus longobardicu
(Reptilia, Protorosauria):
fc-interpretations of thè anatomy based on
■ew specimens from thè Middle Triassic of
■esano (Lombardy, northem Italy)
Memorie Soc. It. Sci Nat. e Museo Civ. St. Nat. Milano
Volume XXXV - Fascicolo III
Piate III - MSNM BES SC 1018
Scale bar 50 mm.
Preparation: Fabio Fogliazza.
Photo: Roberto Appiani.
III
1
NOSOTTI S. - Tanystropheus longobardicus
[(Reptiha, Proiorosauria): Memorie Soc. It. Sci Nat. e Museo Civ. St. Nat. Milano
1
Ili - PELOSIO G., 1968 - Ammoniti del Lias superiore (Toarciano) dell’Al¬
pe Turati (Erba, Como). Generi Hildoceras, Phymatoceras, Paronice-
ras e Frechiella. Conclusioni generali, pp. 143-204, 2 figg. , 6 tavv.
Volume XVIII
I - PINNA G., 1969 - Revisione delle ammoniti figurate da Giuseppe Me¬
neghini nelle Taw. 1-22 della « Monographie des fossiles du calcaire
rouge ammonitique» ( 1 867- 1881). pp. 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 - PETRUCCI F., BORTOLAMI G. C. & DAL PIAZ G. V., 1970 - Ri¬
cerche sull’anfiteatro morenico di Rivoli-Avigliana (Prov. Torino) e sul
suo substrato cristallino, pp. 93-169, con carta a colorì a! 1:40.000, 14
figg., 4 taw. a colorì e 2 b.n.
Volume XIX
I - CANTALUPPI G., 1970 - Le Hildoceratidae del Lias medio delle regio¬
ni mediterranee - Loro successione e modificazioni nel tempo. Riflessi
biostratigrafici e sistematici, pp. 5-46, 2 tabb. n.t.
II - PINNA G. & LEVI-SETTI F., 1971 - I Dactylioceratidae della Provin¬
cia Mediterranea (Cephalopoda Ammonoidea). pp. 47-136, 21 figg.,
12 taw.
Ili - PELOSIO G., 1973 - Le ammoniti del Trias medio di Asklepieion (Ar-
golide, Grecia) - I. Fauna del «calcare a Ptychites » (Anisico sup.). pp.
137-168, 3 figg., 9 taw.
Volume XX
I - CORNAGGIA CASTIGLIONI O., 1971 - La cultura di Remedello.
Problematica ed ergologia di una facies dell’Eneolitico Padano, pp.
5-80, 2 figg., 20 taw.
II - PETRUCCI F., 1972 - Il bacino del Torrente Cinghio (Prov. Parma).
Studio sulla stabilità dei versanti e conservazione del suolo, pp. 81-127,
37 figg., 6 carte tematiche.
Ili - CERETTI E. & POLUZZI A., 1973 - Briozoi della biocalcarenite del
Fosso di S. Spirito (Chieti, Abruzzi), pp. 129-169, 18 figg., 2 taw.
Volume XXI
I - PINNA G., 1 974 - 1 crostacei della fauna triassica di Cene in Val Seriana
(Bergamo), pp. 5-34, 16 figg., 16 taw.
II - POLUZZI A., 1975 - 1 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./?/?. 79-128, 4 figg., 10 taw.
Volume XXII
I - CORNAGGIA CASTIGLIONI O. & CALEGAR1 G„ 1978 - Corpus
delle pintaderas preistoriche italiane. Problematica, schede, iconogra¬
fia. pp. 5-30, 6 figg., 13 taw.
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 taw.
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„ 1982-1 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 taw.
II - POLUZZI A., 1982 - I Radiolari quaternari di un ambiente idrotermale
del Mar Tirreno, pp. 45-72, 3 figg., 1 lab., 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., 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./?/?. 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,
575 figg.
Volume XXVI
I - PINNA G., 1992 - Cyamodus hildegardis Peyer, 1931 (Reptilia, Placo-
dontia). pp. 1-21, 23 figg.
II - CALEGARI G. a cura di, 1993 - L’arte e l’ambiente del Sahara preisto¬
rico: dati e interpretazioni./?/?. 25-556, 647 figg.
III - ANDRI E. e ROSSI F., 1993 - Genesi ed evoluzione di frangenti,
cinture, barriere ed atolli. Dalle stromatoliti alle comunità di scogliera
moderne./?/?. 559-610, 49 figg., 1 tav.
Volume XXVII
I - PINNA G. and GHISELIN M. edited by, 1996 - Biology as History. N.
1. Systematic Biology as an Historical Science./?/?. 1-133, 68 figs.
II - LEONARDI C. e SASSI D. a cura di, 1997 - Studi geobotanici ed en-
tomofaunistici nel Parco Regionale del Monte Barro, pp. 135-266, 122
figg., 23 tabb.
Volume XXVIII
I - BANFI E. & GALASSO G., 1998 - La flora spontanea della città di
Milano alle soglie del terzo millennio e i suoi cambiamenti a partire dal
1700 .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 taw.
Ili - NOSOTTI S. & RIEPPEL O., 2003 - Eusaurosphargis dalsassoi n. gen.
n. sp., a new, unusual diapsid reptile from thè Middle Triassic of Besano
(Lombardy, N Italy)./?/?. 1-33, 19 figs., 1 tab., 3 pls.
Volume XXXII
I - ALESSANDRELLO A., BRACCHI G. & RIOU B., 2004 - Polychaete,
sipunculan and enteropneust worms from thè Lower Callovian (Middle
Jurassic) of La Voulte-sur-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.
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). 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. 1-80, 7 figs., 1 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.
Volume XXXIV
I - GARASSINO A. & SCHWEIGERT G„ 2006 - The Upper Jurassic
Solnhofen decapod crustacean fauna: review of thè types from old
descriptions. Part I. Infraorders Astacidea, Thalassinidea and Palinura.
pp. 1-64, 12 figs., 20 pls.
II - FUCHS D., 2006 - Morphology, taxonomy and diversity of vampyropod
Coleoids (Cephalopoda) from thè Upper Cretaceous of Lebanon. pp.
1-28, 9 figs., 9 pls.
Ili - CALDWELL M. W., 2006 - A new species of Pontosaurus (Squamata,
Pythonomorpha) from thè Upper Cretaceous of Lebanon and a phylo-
genetic analysis of Pythonomorpha. pp. 1-42, 18 figs., 1 pi.
Volume XXXV
I - DE ANGELI A. & GARASSINO A., 2006 - Catalog and bibliography of
thè fossil Stomatopoda and Decapoda from Italy. pp. 1-95.
II - GARASSINO A., FELDMANN R.M. & TERUZZI G„ edited by, 2007 -
3rd Symposium on Mesozoic and Cenozoic Decapod Crustaceans.
Museo di Storia Naturale di Milano May 23-25, 2007. pp. 1-104, 38
figs., 6 tabs.
Le Memorie sono disponibili presso la Segreteria della Società Italiana di Scienze Naturali,
Museo Civico di Storia Naturale, Corso Venezia 55 -20121 Milano
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