© Author(s) 2020. CC Atribution 4.0 License
A new xanthid crab, Neoliomera zovoensis sp. nov. (Decapoda, 
Brachyura), from the Lower Eocene beds of Zovo  
(Vestenanova, Verona, northeast Italy) 
Neoliomera zovoensis sp. nov. (Decapoda, Brachyura), nova vrsta rakovice  
iz spodnjeeocenskih plasti nahajališča Zovo  
(Vestenanova, Verona, severovzhodna Italija) 
Antonio DE ANGELI1 & Alessandro GARASSINO2
1Museo Civico “G. Zannato”, Montecchio Maggiore, Vicenza, Italia; antonio.deangeli@alice.it
2Research Adjunct, Department of Earth and Biological Sciences, Loma Linda University, Loma Linda,  
California, USA; e-mail: alegarassino@gmail.com
Prejeto / Received 4. 11. 2019; Sprejeto / Accepted 2. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020
Key words: Crustacea, Xanthidae, taxonomy, Paleogene, Italy
Ključne besede: Crustacea, Xanthidae, taksonomija, paleogen, Italija
Abstract
A new species of Neoliomera Odhner, 1925, Neoliomera zovoensis sp. nov. from the Lower Eocene (Ypresian) 
of Zovo (Vestenanova, Verona), which co-occurs with other decapod crustacean species in a richly fossiliferous 
coral-algal-reef in the Monti Lessini (Verona area, northeast Italy) is herein described.
Izvleček
V prispevku opisujemo novo vrsto rodu Neoliomera Odhner, 1925, Neoliomera zovoensis sp. nov. iz spodnjega 
eocena (Ipresij) iz nahajališča v Zovu (Vestenanova, Verona). Nova vrsta se pojavlja v združbi z drugimi 
deseteronožci v fosilno bogatem grebenskem apnencu s koralami in algami v regiji Monti Lessini (območje 
Verone, severovzhodna Italija).
GEOLOGIJA 63/1, 67-72, Ljubljana 2020
https://doi.org/10.5474/geologija.2020.007
Introduction
The rich arthropod fauna from the Paleogene 
levels in the Veneto region, which includes mys-
idaceans, isopods, stomatopods, and decapod 
crustaceans, has been recorded over the last two 
centuries. The decapod crustacean assemblage 
is rich in genera and species (see Fabiani, 1910; 
De Angeli & Beschin, 2001). Recent fieldwork has 
yielded numerous new decapod species in asso-
ciation with a bioherm or a small sized coral reef 
(for a checklist of species and complete referenc-
es see De Angeli & Garassino, 2006; De Angeli et 
al., 2019).
Geological and stratigraphical setting
The Bolca area (Verona, northeast Italy; Fig. 1) 
with fossil-rich deposits of “Pesciara” and Monte 
Postale, is renowned for the exceptional preser-
vation of plants, invertebrates, and vertebrates, 
chiefly fishes. For a detailed description of the lo-
cal geology and stratigraphy, see Fabiani (1914, 
1915), Barbieri & Medizza (1969), Medizza (1980a, 
b), and Pasini et al. (2019). 
Above the Scaglia Rossa (Late Cretaceous, 
Campanian) follows the so-called “Calcari di 
Spilecco” (late Paleocene-early Eocene), which is 
in turn overlain by Lithothamnium and Nummu-
lites limestones, the fish-bearing strata of Pescara 
and Monte Postale, and the Alveolina limestones, 
plus marine, brackish, and terrestrial limestones 
of Monte Postale. Higher upsection, Alveolina 
and Nummulites limestones are exposed (hamlet 
of Brusaferri), a thick volcanic mass, containing 
terrestrial plants and freshwater molluscs (Mon-
te Vegroni), shales with Cypris ostracod shells, 
and a coal bed with crocodilian and turtle re-
mains (Monte Purga). The uppermost unit, at the 
68 Antonio DE ANGELI & Alessandro GARASSINO
top of Monte Purga, comprises columnar basalts. 
The uppermost stratified limestones along the 
northern side of Purga di Bolca are dated as late 
Ypresian (Barbieri & Medizza, 1969). The age of 
the reptile-bearing coal beds is still uncertain, 
but could possibly be Lutetian (middle Eocene).
The studied specimen was collected from 
white crystalline limestones with alveolinid 
and nummulitid foraminifera (Brusaferri Lime-
stones), which also contain volcanic material, to 
the southeast of Bolca, along the road connecting 
Zovo and the hamlet of Vallecco (Fig. 1). The fos-
siliferous level at Zovo, containing coralligenous 
algae, corals, micro foraminifera, scarce mollus-
can internal moulds, and exuviae of small- sized 
decapod crustaceans, was associated with a bi-
oherm or a small- sized coral reef, which origi-
nated within the Alpone Agno graben. Forma-
tions such as this have been recorded from the 
Valle del Chiampo between Mussolino and Zovo 
di Castelvecchio (De Zanche, 1965) and along the 
eastern margin of Monti Lessini (Beschin et al., 
2007; De Angeli & Garassino, 2002; De Angeli & 
Ceccon, 2012). Currently, the decapod crustacean 
assemblage from Zovo includes 24 species (for 
complete references see Beschin et al. 2016; De 
Angeli et al., 2019).
Material 
One carapace preserving its entire cuticle 
within a small piece of coralligenous rock. It is 
housed in the palaeontological collection of the 
Museo Civico “D. Dal Lago” of Valdagno, Vicen-
za (MCV).
Abbreviations – lcxp: carapace length, wcxp: 
carapace width, wf: front width, wof: orbito-fron-
tal width;
Systematic palaeontology
For the higher-level classification, we follow 
the arrangement proposed by Schweitzer et al. 
(2010).
Order Decapoda Latreille, 1802
Infraorder Brachyura Latreille, 1802
Section Eubrachyura de Saint Laurent, 1980
Subsection Heterotremata Guinot, 1977
Superfamily Xanthoidea MacLeay, 1838
Family Xanthidae MacLeay, 1838
Subfamily Liomerinae Sakai, 1976
Genus Neoliomera Odhner, 1925
Type species: Zozymus pubescens H.Milne 
Edwards, 1834, by original designation.
Included fossil species: Neoliomera interme-
Fig. 1. Map of the Bolca 
area; the asterisk denotes 
the locality where the type 
specimen of Neoliomera 
zovoensis sp. nov. was co-
llected (modified after De 
Angeli & Garassino, 2014).
69A new xanthid crab, Neoliomera zovoensis sp. nov. (Decapoda, Brachyura), from the Lower Eocene beds of Zovo...
dia Odhner, 1925 (fossil and extant), N. kuohwai 
Hu, 1981 (fossil), N. minuta Beschin, Busulini 
& Tessier, 2015 (fossil), N. paleogenica Beschin, 
Busulini, De Angeli & Tessier, 2007 (fossil), N. 
pubescens (H. Milne Edwards, 1834) (fossil and 
extant), N. richteroides Sakai, 1969 (fossil and 
extant); N. zovoensis sp. nov. (herein).
Neoliomera zovoensis sp. nov.
(Figs. 2.1a, 1b)
Diagnosis: Subhexagonal carapace, convex 
longitudinally, broader than long; bilobed front; 
small suboval orbits; raised, granulate supraor-
bital margin; elongate, convex anterolateral 
Fig. 2. Neoliomera zovoensis sp. nov., Holotype, MCV 19/09. 1a - carapace in frontal view; 1b - carapace in dorsal view. Scale 
bar equals 5 mm.
70 Antonio DE ANGELI & Alessandro GARASSINO
margins, with four short spiny lobes; fourth an-
terolateral lobe with granulate ridge extending 
on branchial region; short, convergent postero-
lateral margins; undifferentiated dorsal regions; 
one longitudinal median groove in frontal region; 
cervical groove dividing hepatic region from 
branchial ones; smooth dorsal carapace surface, 
except for some tubercles uniformly arranged in 
frontal region and in the outer parts of hepatic 
and epibranchial regions.
Etymology: after Zovo where the studied 
specimen was discovered.
Holotype: MCV 19/09.
Type locality: Zovo (Vestenanova, Verona, 
northeast Italy).
Measurements: MCV 19/09 – lcxp: 13.3 mm; 
wcxp: 29.5 mm; wof: 11.5 mm; wf: 8 mm.
Description: Subhexagonal carapace, convex 
longitudinally, broader than long (lcxp/wcxp 
= 0.45); orbito-frontal margin moderately wide 
(wof/wcxp = 0.38); bilobed front grooved medi-
ally and downturned; frontal margin with small 
tubercles arranged uniformly; small subround 
orbits with raised, granulate supraorbital mar-
gin; convex inner orbital angle well distinct from 
the front by an indentation; elongate, convex 
anterolateral margins, with two or three small 
spines, close each other, forming four short con-
vex spiny lobes, divided by weak fissures: first 
with two small spines (excluding the extra-orbit-
al tooth), second and third with three spines, and 
fourth, at level of anterolateral angle, with one 
spine and one granulate ridge (branchial ridge) 
extending on the branchial region; shorter pos-
terolateral margins, strongly converging to the 
posterior margin; posterior margin as wide as the 
front, weakly convex and rimmed; undifferenti-
ated dorsal regions; one deep longitudinal medi-
an groove in the frontal region; cervical groove 
dividing hepatic region from the branchial ones; 
weak transverse depression in the cardiac region; 
weak branchiocardiac grooves, more evident 
along the cardiac region margins; smooth dor-
sal carapace surface, except some tubercles uni-
formly arranged in the frontal region and in the 
outer parts of hepatic and epibranchial regions; 
small pits arranged uniformly on dorsal surface, 
richer in frontal and hepatic regions. 
Discussion: Based upon Sakai (1976) and 
Serène (1984), the studied specimen shows the 
main morphological characters of the extant 
Neoliomera in having a carapace broader than 
long; anterolateral margins crested and entire, 
although marked with three to four demarcated, 
rounded lobes; and poorly defined regions. Neoli-
omera is currently widely distributed in the In-
do-West Pacific area with 17 species inhabiting 
rocky beach, under stones or in coral reef, and 
shallow waters (Ho & Ng, 2014). 
Neoliomera is known in the fossil record of 
northern Italy with two species, N. paleogenica 
Beschin, Busulini, De Angeli & Tessier, 2007, 
from the early Eocene of contrada Gecchelina 
(Monte di Malo, Vicenza) and N. minuta Beschin, 
Busulini & Tessier, 2015 from the early Eocene 
of Cava Braggi (Vestenanova, Verona). The for-
mer differs from N. zovoensis sp. nov. in having 
meso-, metagastric, and cardiac regions that are 
well differentiated by grooves and thick tuber-
culate ornamentation uniformly arranged on the 
whole dorsal surface (Beschin et al., 2007). The 
latter differs from the new species in having a 
more oval carapace outline, dorsal surface of car-
apace with randomly arranged small tubercles, 
an anterior mesogastric process that does not 
reach the front, a carapace that is not marked by 
a cervical groove, an anterolateral margin with 
four smooth lobes, and an anterolateral angle 
without branchial ridge (Beschin et al., 2015). 
Neoliomera zovoensis sp. nov. has a shallow 
cervical groove and a weak granulate ridge on 
the branchial regions, as in the extant N. themis-
to (De Man, 1889), widespread in the Persian 
Gulf (see Guinot, 1964). This extant species dif-
fers, however, from the fossil one in having more 
distinct hepatic and branchial regions with larg-
er and more numerous tubercles.
Acknowledgements
We wish to thank Bernardetta Pallozzi (Museo 
Civico “D. Dal Lago”, Valdagno, Vicenza) for ma-
king the specimen available for study, and Rodney 
M. Feldmann, (Kent State University, Ohio, USA) 
and Matúš Hyžný (Comenius University, Bratislava, 
Slovakia) for careful review and criticism.
References
Barbieri, G. & Medizza, F. 1969: Contributo alla 
conoscenza geologica della regione di Bolca 
(Monti Lessini). Memorie Istituto Geologico 
Mineralogico dell’Università di Padova, 27: 
1-36.
Beschin, C., Busulini, A., De Angeli, A. & Tessier, 
G. 2007: I Decapodi dell’Eocene inferiore di 
Contrada Gecchelina (Vicenza - Italia setten-
trionale) (Anomura e Brachyura). Museo di 
Archeologia e Scienze Naturali “G. Zannato”, 
Montecchio Maggiore (Vicenza): 76 p.
71A new xanthid crab, Neoliomera zovoensis sp. nov. (Decapoda, Brachyura), from the Lower Eocene beds of Zovo...
Beschin, C., Busulini, A. & Tessier, G. 2015: 
Nuova segnalazione di crostacei associati a 
coralli nell’Eocene inferiore dei Lessini ori-
entali (Vestenanova - Verona). Lavori, Società 
Veneziana di Scienze Naturali, 40: 47-109.
Beschin, C., Busulini, A., Tessier, G. & Zorzin, R. 
2016: I crostacei associati a coralli nell’Eocene 
inferore dell’area di Bolca (Verona e Vicenza, 
Italia nordorientale). Memorie Museo Civico 
di Storia Naturale di Verona, 2. Ser., Sezione 
Scienze della Terra, Verona, 9: 1-190.
De Angeli, A. & Beschin, C. 2001: I Crostacei 
fossili del territorio Vicentino. Natura 
Vicentina, 5: 5-54.
De Angeli, A. & Ceccon, L. 2012: Eouroptychus 
montemagrensis n. gen., n. sp. (Crustacea, 
Decapoda, Anomura, Chirostylidae) 
dell’Eocene inferiore (Ypresiano) di Monte 
Magrè (Vicenza, Italia settentrionale). Società 
Veneziana di Scienze Naturali, 37: 19-24.
De Angeli, A. & Garassino, A. 2002: Galatheid, 
chirostylid and porcellanid decapods 
(Crustacea, Decapoda, Anomura) from the 
Eocene and Oligocene of Vicenza (N Italy). 
Memorie della Società italiana di Scienze 
naturali e del Museo civico di Storia na-
turale di Milano, 30/3: 1-40. https://doi.
org/10.1163/18759866-0720203006
De Angeli, A. & Garassino, A. 2006: Catalog and 
bibliography of the fossil Stomatopoda and 
Decapoda from Italy. Memorie della Società 
italiana di Scienze naturali e del Museo ci-
vico di Storia naturale di Milano, 35/1: 1-95.
De Angeli, A. & Garassino, A. 2014: A new genus 
and species of panopeid crab, Zovocarcinus 
muelleri (Decapoda, Brachyura) from the 
Eocene of Zovo di Bolca, Verona (northeast 
Italy). In: Fraaije, R.H.B., Hyžný, M., Jagt, 
J.W.M., Krobicki M. & Van Bakel, B.W.M. 
(eds.), Proceedings of the 5th Symposium on 
Mesozoic and Cenozoic Decapod Crustaceans, 
Krakow, Poland, 2013: A tribute to Pál Mihály 
Müller. Scripta Geologica, 147: 185-191.
De Angeli, A., Garassino, A. & Pasini, G. 
2019: Catalog and bibliography of fossil 
Stomatopoda and Decapoda from Italy (2007-
2018). Memorie della Società italiana di 
Scienze naturali e del Museo civico di Storia 
naturale di Milano, 45: 3-70.
De Zanche, V. 1965: Le microfacies eoceniche 
nella Valle del Chiampo tra Arzignano e 
Mussolino (Lessini orientali). Rivista Italiana 
di Paleontologia e Stratigrafia, 71/3: 925-948.
Fabiani, R. 1910: I crostacei terziari del Vicentino. 
Bollettino del Museo Civico di Vicenza, 1/1: 1-40.
Fabiani, R. 1914: La serie stratigrafica del Monte 
Bolca e dei suoi dintorni. Memorie dell’Is-
tituto Geologico della Reale Università di 
Padova, 2: 223-235.
Fabiani, R. 1915: Il Paleogene del Veneto. Memorie 
dell’Istituto Geologico della Reale Università 
di Padova, 3: 1-336.
Guinot, D. 1964: Crustacés Décapodes 
Brachyoures (Xanthidae) des Campagnes de 
la Calypso en Mer Rouge (1952), dans le Golfe 
Persique et a l’Ile Aldabra (1954). Mémoires 
du Muséum national d’Histoire naturelle, 
32/1: 1-108.
Guinot, D. 1977: Proposition pour une nouvel-
le classification des Crustacés Décapodes 
Brachyoures. Comptes Rendus hebdomadai-
res des Séances de l’Académie des Sciences, 
(D), 285: 1049-1052. 
Ho, P.-H. & Ng, P.K.L. 2014: A new species of 
Neoliomera Odhner, 1925, from the western 
Pacific, and the first record of N. demani Forest 
& Guinot, 1961, from Taiwan (Crustacea: 
Decapoda: Brachyura: Xanthidae). Zootaxa, 
3826/3: 579-590. https://doi.org/10.11646/
zootaxa.3826.3.8
Hu, C.-H. 1981: Studies on Cenozoic fossil crabs 
from Taiwan Island. Proceedings of the 
Geological Society of China, 24: 56-74. 
Latreille, P.A. 1802-1803: Histoire naturelle 
générale et particulière, des Crustacés et des 
Insectes. Ouvrage faisant suite aux uvres de 
Leclerc de Buffon, et partie du Cours complet 
d’Histoire naturalle rédigé par C.S. Sonnini, 
membre de plusieurs Société savantes. 5: 
1-407, 6: 1-391; Paris (Dufart). 
MacLeay, W.S. 1838: On the Brachyurous 
Decapod Crustacea. Brought from the Cape 
by Dr. Smith. Illustrations of the Zoology of 
South Africa; consisting chiefly of figures 
and descriptions of the objects of natural his-
tory collected during an expedition into the 
interior of South Africa, in the years 1834, 
1835, and 1836; fitted out by “The Cape of 
Good Hope Association for Exploring Central 
Africa:” together with a summary of African 
Zoology, and an inquiry into the geographical 
ranges of species in that quarter of the globe, 
published under the Authority of the Lords 
Commissioners of Her Majesty’s Treasury, 
Invertebratae. IV (1849), London (Smith, 
Elder & Co.): 53-71. 
Man, J. G. de 1889: Über einige neue oder selte-
ne indo-pacifische Brachyuren. Zoologische 
Jahrbücher, Iena, (Systematik) 4: 409-452. 
https://doi.org/10.5962/bhl.part.18727
72 Antonio DE ANGELI & Alessandro GARASSINO
Medizza, F. 1980a: Il giacimento di Spilecco 
(Verona). In: I vertebrati fossili italiani. 
Catalogo della Mostra, Verona, 139-141.
Medizza, F. 1980b: Il giacimento della Purga di 
Bolca (Verona). In: I vertebrati fossili italiani. 
Catalogo della Mostra, Verona, 147-148.
Milne Edwards, H. 1834-1837: Histoire naturel-
le des Crustacés comprenant l’anatomie, la 
physiologie et la classification de ces anima-
ux. Paris. Librairie Encyclopédique de Roret. 
1: 1-468, 2: 1-531, 3: 1-638.
Odhner, T. 1925: Monographierte Gattungen der 
Krabbenfamilie Xanthidae. I. Göteborgs 
Kungliga Vetenskaps-och Vitterhets-
Samhälles Handlingar, ser. 4, 29/1: 1-92. 
Pasini, G., Garassino, A., De Angeli, A., Hyžný, M., 
Giusberti, L. & Zorzin, R. 2019: Eocene deca-
pod faunas from the Konservat-Lagerstätten 
laminites of “Pesciara” (Bolca, Verona) 
and Monte Postale (Altissimo, Vicenza) in 
northeast Italy: a review and update. Neues 
Jahrbuch für Geologie und Paläontologie, 
Abhandlungen 293/3: 233-270. https://doi.
org/10.1127/njgpa/2019/0840 
Sakai, T. 1969: Two new genera and twenty-two 
new species of crabs from Japan. Proceedings 
of the Biological Society of Washington, 82: 
243-280. 
Sakai, T. 1976: Crabs of Japan and the Adjacent 
Seas. In three volumes; Kodansha Ltd., Tokyo; 
English Text, 773 pp., Japanese Text, 461 pp. 
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 heb-
domadaires des Séances de l’Académie des 
Sciences, (D), 290: 1265-1268. 
Serène, R. 1984: Crustacés Dédapodes 
Brachyoures de l’Océan Indien Occidentale 
et de la Mer Rouge. Xanthoidea: Xanthidae 
et Trapeziidae. Addendum Carpiliidae et 
Menippidae. A. Crosnier. Faune Tropicale, 24: 
1-400.
Schweitzer, C.E., Feldmann, R.M., Garassino, 
A., Karasawa, H. & Schweigert, G. 2010: 
Systematic list of fossil decapod crustacean 
species. Crustaceana Monographs, 10: 1-222.