Lophoranina maxima Beschin, Busulini, De Angeli & Tessier, 2004 (Decapoda Brachyura, Raninidae) from Lower Eocene laminites of the "Pesciara di ...
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GEOLOGIJA 63/1, 57-66, Ljubljana 2020 © Author(s) 2020. CC Atribution 4.0 License
https://doi.org/10.5474/geologija.2020.006
Lophoranina maxima Beschin, Busulini, De Angeli & Tessier,
2004 (Decapoda Brachyura, Raninidae) from Lower Eocene
laminites of the “Pesciara di Bolca” (Verona, northeast Italy)
Lophoranina maxima Beschin, Busulini, De Angeli & Tessier, 2004 (Decapoda
Brachyura, Raninidae) iz spodnjeeocenskih laminiranih apnencev “Pesciara di Bolca”
(Verona, severovzhodna Italija)
Alessandra BUSULINI1, Roberto ZORZIN2, Claudio BESCHIN3 & Giuliano TESSIER1
1
Società Veneziana di Scienze Naturali, Museo di Storia Naturale “Giancarlo Ligabue”, Santa Croce 1730, 30135
Venezia, Italy; e-mail: busulini@tin.it; giultess@virgilio.it
2
Museo Civico di Storia Naturale di Verona, Lungadige Porta Vittoria 9, 37129 Verona, Italy;
e-mail: roberto.zorzin@comune.verona.it
3
Museo Civico “G. Zannato”, Piazza Marconi 15, 36075 Montecchio Maggiore (Vicenza), Italy;
e-mail: beschin.cl@libero.it
Prejeto / Received 1. 12. 2019; Sprejeto / Accepted 7. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020
Key words: Crustacea, Decapoda, taphonomy, Paleogene, Mediterranean
Ključne besede: raki, deseteronožci, tafonomija, paleogen, Sredozemlje
Abstract
The sole specimen of a raninid crab found to date in the Lower Eocene Fossil-Lagerstätte of the “Pesciara di
Bolca” (Verona, northeast Italy) and referred to Lophoranina maxima, is described. Results of a CT analysis of
this specimen and of a study of its cuticle are discussed.
Izvleček
Opisan je edini primerek raninidne rakovice, ki je bila do sedaj najdena v spodnjeeocenskih apnencih
znanega nahajališča "Pesciara di Bolca" (Verona, severovzhodna Italija) in pripada vrsti Lophoranina maxima.
Predstavljeni so rezultati CT analize primerka in analiza kutikule oklepa.
Introduction
Ocypodidae Rafinesque, 1815. Since then some of
Among representatives of the so-called “minor these remains have been revised and additional
fauna” from the Lower Eocene Fossil-Lagerstätte specimens recovered. Vonk et al. (2015) studied
called “La Pesciara” (Bolca, Vestenanova, Vero- the Isopoda, while additional papers dealt with
na, northeast Italy; Fig. 1) abundant specimens stomatopods (see Giusberti et al., 2014). Final-
of malacostracan crustaceans have been found; ly, Pasini et al. (2019a, b) thoroughly revised the
these are referred to the Isopoda, Decapoda and crustacean fauna from this area.
Stomatopoda. Most of these are housed in the The sole specimen of the genus Lophorani-
collections of the Natural History Museum in na Fabiani, 1910a from the “Pesciara” laminites
Verona, the Fossil Museum at Bolca, the Univer- (Fig. 2) is on display in the permanent exhibition
sity of Padua and the Natural History Museum of the Natural History Museum in Verona. De-
in Milan. After incomplete analyses published spite easy access to this fossil it has never been
during the 19th century, they were studied in de- analysed in full. The reason why it was not includ-
tail by Secretan (1975). Amongst brachyurans ed in her study by Secretan (1975) is unknown:
she described species of the families Macropip- at that time, it certainly was already housed in
idae Stephenson & Campbell, 1960, Portunidae the palaeontological collections of the Veronese
Rafinesque, 1815, Panopeidae Ortmann, 1893, museum. It was recorded for the first time by
Eriphiidae MacLeay, 1838 and, probably, of the Beschin et al. (2011) as Lophoranina marestiana
58 Alessandra BUSULINI, Roberto ZORZIN, Claudio BESCHIN & Giuliano TESSIER
level near the watershed between the high Al-
pone Valley (Vestenanova, Verona) and the Chi-
ampo Valley (Altissimo, Vicenza). This territory
is part of the Southern Alpine tectonic unit; in
a general geological framework it is constitut-
ed mainly by volcanic rocks and secondarily by
small outcrops of carbonate rocks of Cretaceous–
Paleogene age that rest on the so-called Trento
Platform palaeogeographical unit. During the
Middle Jurassic it formed a structural high be-
tween the Lombard and Belluno lateral basins;
later it completely subsided and up to the Pale-
ocene it assumed the character of a pelagic pla-
teau with marine sedimentation. During Alpine
Orogeny this area responsed rigidly to tectonic
stresses and was broken up into blocks. Some of
these rose up to shallow sea conditions; on these
small carbonate platforms came into existence.
Afterwards they built a unique structure called
the Lessini Shelf (Bosellini, 1989).
Fig. 1. Location of the “Pesciara di Bolca” site (modified af-
ter Beschin et al., 2016b). Important faults produced during the Pale-
ocene and Early and Middle Eocene activated
(König, 1825), but not described. Later, Zorzin et volcanic cycles with emissions of great amounts
al. (2019) referred it to Lophoranina maxima and of basic lavas, mostly submarine, associated with
this attribution has subsequently been confirmed extensional tectonics (Piccoli, 1965, 1966). Late
by Pasini et al. (2019b). This note presents a de- Eocene volcano-tectonic activity resulted in the
tailed description and analysis of this particular opening of the Alpone-Agno graben or semigra-
individual. ben, a wide and lengthy depression delimited to
the west by the Castelvero Fault and to the east
Geological setting of the by the Schio-Vicenza fault system. During quies-
“Pesciara di Bolca” cent phases of volcanism in the Early and Middle
The Bolca area, with its famous Fossil-La- Eocene, carbonate sediments and marls were cy-
gerstätten of the “Pesciara” and Monte Postale, is clically laid down in this graben (Barbieri et al.,
located in the eastern part of the Lessini Moun- 1991; Zampieri, 1995). Despite intense magmatic
tains at an altitude of about 800 metres above sea activity, a rich fauna, represented by crustaceans,
Fig. 2. Panoramic view of
the “Pesciara” site.
Lophoranina maxima Beschin, Busulini, De Angeli & Tessier, 2004 (Brachyura, Raninidae) from Lower Eocene laminites... 59
bivalves, gastropods, echinoids and other macro- tions of the Natural History Museum in Verona
biota inhabited the sea floor. Fossils are perfectly (Museo Civico di Storia Naturale di Verona), un-
preserved and particularly abundant within the der registration numbers CR 55 and CR 56.
Eocene tuffites cropping out in the Chiampo Val- CR 55 is the negative (Figs. 3a, b), CR 56 the
ley and in the Roncà area (central and southern positive (Figs. 4a, b). CR 56 was computed tomog-
Alpone Valley): they have been studied with great raphy (CT) scanned and its cuticle was studied
interest since the 18th century. with the use of a stereoscopic microscope (Leica
Tectonic stresses produced fragile respons- M165C).
es on the calcareous rocks with the isolation of a
series of blocks bounded by faults; some of these
collapsed as olistolithes in the basin (Barbieri & Systematic palaeontology
Zampieri, 1992; Zampieri, 1995). This is the case
for the “Pesciara” outcrop which is made up of a Order Decapoda Latreille, 1802
series of Lower Eocene calcareous strata, about 19 Infraorder Brachyura Linnaeus, 1758
metres thick, and completely surrounded by vol- Section Podotremata Guinot, 1977
canoclastic rocks (Papazzoni & Trevisani, 2006). Subsection Gymnopleura Bourne, 1922
Fossils recovered from the “Pesciara”, mostly Superfamily Raninoidea De Haan, 1839
fish and plants, are preserved in five successive Family Raninidae De Haan, 1839
horizons made of extremely fine-grained, thinly Genus Lophoranina Fabiani, 1910a
laminated limestones that are interbedded with
detrital calcareous levels that yield invertebrate Type species: Ranina marestiana König, 1825,
remains, mostly larger foraminifera, bivalves and by original designation.
gastropods. This succession of fine- and coarse-
grained limestones testifies to cyclic phases of Lophoranina maxima Beschin, Busulini,
different environmental conditions. De Angeli & Tessier, 2004
The environment of the “Pesciara” is charac- 1983 Lophoranina reussi; Busulini et al., p. 61,
terised by deposition of calcareous muds within pl. 2, fig. 1 (non Woodward, 1866).
an intra-platform basin, in which anoxic bottom 1988 Lophoranina cf. reussi; Beschin et al., p.
conditions and a microbial film that developed on 185, fig. 8, pl. 5, fig. 1; pl. 8, figs 1-4; pl. 9, fig. 1.
corpses enabled perfect preservation of its rich 2004 Lophoranina maxima Beschin, Busulini,
and diverse fossil fauna (Marramà et al., 2016; De Angeli & Tessier, p. 110, text-figs. 1, 2; pl. 1,
Friedman & Carnevale, 2018). The presence of figs. 1-3; pl. 2, figs. 1, 2.
coral reefs that are similar to the ones observed 2006 Lophoranina maxima; De Angeli & Ga-
along the coasts of the present-day St. Croix Is- rassino, p. 35.
land (Caribbean Sea) cannot be excluded, at least 2010 Lophoranina maxima; Schweitzer et al.,
along the outer margin of the “Pesciara” lagoon p. 73.
(Beschin et al., 2017). 2011 Lophoranina marestiana; Beschin et al.,
The “Pesciara” is the most famous and im- p. 38 (pars).
portant Eocene Fossil-Lagerstätte in Italy. Ex- 2011 Lophoranina maxima; Beschin et al., p.
cavations since the 16th century, and particularly 46, text-fig. 9, pl. 4, fig. 1.
those in the 2000s on the instigation of the Muse- 2019 Lophoranina maxima; Zorzin et al., p.
um of Natural History in Verona, have allowed to 97, figs. 1, 2.
recover not only a great number of fish (Bosellini 2019b Lophoranina maxima; Pasini et al., p.
et al., 2014), but also remains of reptiles and birds 261, fig. 17A, B.
and a rich “minor fauna”. The latter comprises
crustaceans (malacostracans and ostracods), Measurements (in mm): Carapace: maximum
scorpions, bivalves, cephalopods, gastropods, width ~ 95; posterior width ~ 50; length > 85.
brachiopods, bryozoans, worms, corals, jellyfish Right carpus: height ~ 20. Right propodus: max-
and foraminifera. imum length ~ 50; maximum height ~ 35. Right
dactylus: length ~ 30; height ~ 10.
Material and methods Description (based on both positive and neg-
The specimen studied here exposes the cara- ative): Subovate carapace, dorsoventrally com-
pace in dorsal view as well as both chelipeds and pressed and strongly damaged in both anterior
is preserved as two slabs (width about 245 mm; and posterior parts; only right part of wide fron-
length about 195 mm); it is housed in the collec- to-orbital margin preserved with a strong and
60 Alessandra BUSULINI, Roberto ZORZIN, Claudio BESCHIN & Giuliano TESSIER
a b
Fig. 3. CR 55, negative (maximum width 95 mm), in photograph (a) and line drawing (b).
a b
Fig. 4. CR 56, positive part (maximum width 95 mm), in photograph (a) and line drawing (b).
pointed lateral tooth and a supraorbital tooth de- the frontal area was short. On terraces, bases of
fined by two fissures (Fig. 5a). Lateral margins small spines that constituted them are visible as
convergent; almost completely preserved and regularly spaced, tiny pits.
appearing double because of separation of upper In median anterior part, there is a structure
and lower part of carapace during fossilisation recognised as the sternal plate using CT scan-
(Fig. 5b). In anterior part of right margin, traces ning (Fig. 6b); it shows some weak transverse
of two large teeth visible; two spiny large teeth terraces that probably are traces of ornament of
can be observed on left margin (Fig. 5c) linked carapace impressed on it.
to a fragment of carapace that was thrown for- Both chelipeds are preserved: left one is set-
wards probably during fossilisation and so dis- tled near carapace, almost in anatomical position
located from original position. Rear part of right and covered with matrix; right one outstretched
lateral margin showing granulated rim (Fig. 5d). and shows propodus and carpus; both large,
Posterior margin, that should have been narrow- stout, covered with subparallel transverse ridg-
er than fronto-orbital margin, is lacking. Dorsal es. Carpus appears almost squarish with a spine
ornament with at least 21 subparallel transverse on upper distal angle; propodus shows a spine on
terraces, well preserved mainly in intermedi- upper margin and three spines on lower margin
ate part of carapace, where terraces are nearly (including fixed finger); dactylus is long and sick-
continuous from one side to the other, while rear le-shaped. Distal part of right fifth pereiopod
ones appear interrupted. As far as can be seen, with paddle-like dactylus.
Lophoranina maxima Beschin, Busulini, De Angeli & Tessier, 2004 (Brachyura, Raninidae) from Lower Eocene laminites... 61
b
a
c d
Fig. 5. Details of carapace: A. CR 55 (negative), right orbital and lateral margins (scale bar equals 10.0 mm); B. CR 56, double
left posterolateral margin (scale bar equals 2.8 mm); C. CR 56, left anterolateral tooth (scale bar equals 2.5 mm); D. CR 56,
right posterolateral margin (scale bar equals 4.0 mm).
Distribution: Lophoranina maxima has previ- composed of tiny, forwardly inclined spines on
ously been recorded only from Lutetian (Middle the dorsal carapace surface. This extinct genus
Eocene) rocks at Main Quarry (Arzignano-Vi- has a worldwide distribution and a stratigraph-
cenza) and Case Pozza di San Giovanni Ilarione ical range from the Eocene to the Miocene. In
(Verona). It is now recognised in Ypresian (Lower Veneto (northeast Italy), representatives of Lo-
Eocene) levels in the “Pesciara di Bolca” (Vero- phoranina are highly characteristic of Eocene
na) and, according to Pasini et al. (2019b), also at levels of volcanoclastic origin and include nu-
Monte Postale. merous species such as L. avesana (Bittner, 1883),
L. bittneri (Lőrenthey, 1902), L. laevifrons (Bit-
Discussion tner, 1875), L. marestiana (König, 1825), L. max-
Fabiani (1910a; see also Fabiani, 1910b) erect- ima, L. reussi (Woodward, 1866), L. straeleni Vía
ed the genus Lophoranina to accommodate spe- Boada, 1959 and, probably, L. aldrovandii (Ran-
cies that had previously been included in Ranina zani, 1818) (see Beschin et al., 1988, 2011, 2016a).
Lamarck, 1801, but showed transverse terraces The structure of the dorsal terraces and the large
62 Alessandra BUSULINI, Roberto ZORZIN, Claudio BESCHIN & Giuliano TESSIER
a that are situated more anteriorly, the propodus of
the cheliped with six spines (including the fixed
finger) on the lower margin. The differences with
the other species are greater.
Our CT analysis of specimen CR 56 (positive)
has revealed that the matrix piece in which the
specimen is preserved was broken into four main
pieces along three straight fractures (Fig. 6a).
One fracture cuts the specimen in the rear part
of the carapace and a second one runs through
the propodus of the right cheliped (Fig. 4a).
During restoration, fragments were fixed with
a thick cement on a calcareous slab (in its turn
divided into two parts) as a reinforcement (total
b
thickness about 37 mm) (Fig. 6c). This prepara-
tion method was applied to fossils found in the
“Pesciara” during the 1930s (Massimo Cipriano
Cerato, pers. comm., 2019; the Cerato family
of Bolca are the owners of the “Pesciara” site,
where excavations have been ongoing for about
two centuries, and during the last fifty years un-
der supervision of the Museum of Natural His-
tory in Verona). Hence, specimens CR 55/66 of
Lophoranina maxima was presumably collected
during those years. The CT axial scan shows that
the frog crab is almost completely dorsoventral-
ly flattened and produced only a weak relief of
c the surface of the slab. The CT coronal recon-
struction shows the outline of the carapace with
a thick fracture in its rear part and the collapsed
cardiac part; a small shield-shaped structure in
the anterior part is reminiscent of the sternal
plate (Fig. 6b); it was probably dislocated during
fossilisation.
The general preservation of this specimen
confirms observations made by Secretan (1975),
Fig. 6. CR 56. A. CT coronal reconstruction of the slab under
the specimen (depth 20.1 mm); B. CT coronal reconstructi- who noted that the crustaceans found in the
on showing the specimen (depth 8.6 mm); C. CT axial scan “Pesciara” were almost flattened and lost any
(81 mm from the anterior margin of the slab).
reliefs, their outlines being “confused” and part
of the cuticle removed and dislocated.
carapace size, the short fronto-orbital region, Cuticle is preserved mainly in specimen CR
the position and number (two) of anterolateral 55 (negative): it shows almost the entire cara-
spines, the relatively short propodus of chelipeds pace cuticle in its inner, deeper part (Fig. 5a).
with only three spines on the lower margin, all In specimen CR 56 (positive) only a few shreds
suggest that specimen CR 55/56 belongs to Lo- of the cuticle can be seen. An analysis using a
phoranina maxima. As far as comparisons with stereoscopic microscope detected thick cuticle
other large-sized species of Lophoranina are in a natural cross section along the lateral mar-
concerned, L. avesana has fewer ridges on the gins of the carapace and at the tip of the dacty-
dorsal surface but these are more continuous and lus, and revealed details of the deep structure of
form a general anterior concavity; moreover, the the finger. As can be seen in Figure 7a, the pre-
anterolateral spines are spatulate rather than served cuticle in its upper margin is composed
spiny, and the propodus of the cheliped is longer of a thick endocuticle that shows undulate lam-
with a dentate lower margin. Lophoranina mar- inations in its deep portion; each undulation
estiana has a longer fronto-orbital area, more corresponds to a small globular swelling in the
regular ridges, less acute anterolateral spines amorphous filling of the dactylus surmounted
Lophoranina maxima Beschin, Busulini, De Angeli & Tessier, 2004 (Brachyura, Raninidae) from Lower Eocene laminites... 63
The presence of a crab with burrowing habits
may appear improbable in a palaeoenvironment
that has generally been considered as anoxic,
but many fish and also crustaceans found in the
“Pesciara” were benthic species. Our analysis
also aimed at determining whether or not this
particular specimen was a moult or a corpse.
Most of the individuals of Lophoranina in vol-
canoclastic rocks in Veneto are moults (in open
moult position, or Salter’s position). The par-
ticular preservation of the specimen found in the
“Pesciara” laminites does not allow this to be
a determined; however, the position of the cheli-
peds and the collapsed cardiac region suggest
that it could be a corpse (Bishop, 1986). The good
condition of the lower cuticle layer is in agree-
ment with this hypothesis, although Waugh et
al. (2009a) pointed out that this feature does not
allow to determine with certainty the nature of
a specimen in fossil material.
Acknowledgements
We wish to thank Francesca Rossi, manager of
“Direzione dei Musei” (Comune di Verona), for per-
mission to study a specimen under her care, Roberta
b Salmaso (Museo Civico di Storia Naturale di Verona)
for providing photographs taken with the stereoscopic
microscope, Mario Calvagno for CT scanning of the
specimen and post-processing of images, Massimo
Cipriano Cerato for useful information on the prepa-
ration methods of “Pesciara” fossils. We express our
gratitude to René Fraaije (Oertijdmuseum, Boxtel, the
Netherlands) and Martina Kočová Veselská (Institute
of Geology, Czech Academy of Sciences, Prague,
Czech Republic) for their kind and thoughtful re-
views of an earlier version of the typescript, and to
John Jagt (Natuurhistorisch Museum Maastricht, the
Netherlands) for his careful assistance and linguistic
help. Thanks also to Matteo Calvagno for help with
graphics.
c
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(Slovenia), 17-21 June 2019: 101-102.You can also read
