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Review focus Journal of the Geological Society

Published online May 8, 2018 https://doi.org/10.1144/jgs2017-164 | Vol. 175 | 2018 | pp. 569–579

The Bolca Lagerstätten: shallow marine life in the Eocene

Matt Friedman1* & Giorgio Carnevale2
1
Museum of Paleontology and Department of Earth and Environmental Sciences, University of Michigan, 1109 Geddes Ave,
Ann Arbor, MI 48109-1079, USA
2
Dipartimento di Scienze della Terra, Università degli Studi di Torino, via Valperga Caluso 35, 10125, Torino, Italy
M.F., 0000-0002-0114-7384
* Correspondence: [email protected]

Abstract: The Eocene limestones around the Italian village of Bolca occur in a series of distinct localities providing a unique
snapshot of marine life in the early Cenozoic. Famous for its fishes, the localities of Bolca also yield diverse invertebrate faunas
and a rich, but relatively understudied flora. Most fossils from Bolca derive from the Pesciara and Monte Postale sites, which
bear similar fossils but are characterized by slightly different taphonomic and environmental profiles. Although not precisely
contemporaneous, the age of these principal localities is well constrained to a narrow interval within the Ypresian Stage, c. 50–
49 Ma. This places Bolca at a critical time in the evolutionary assembly of modern marine fish diversity and of reef
communities more generally.
Received 22 December 2017; revised 7 March 2018; accepted 8 March 2018

The rich fossil sites near Bolca, Italy provide a picture of life in a contains remains of crocodiles, turtles, snakes and plants. The
warm, shallow marine setting during the early Eocene, roughly lignites of Vegroni yield a variety of plants.
50 myr ago. Deposited within an archipelago near the western end
of the Tethys Ocean, the major localities of Bolca collectively have
yielded over 500 species of terrestrial vertebrates, fishes, insects, Stratigraphy and age
marine invertebrates and plants. The outstanding significance of the
The Pesciara and Monte Postale successions are among the few
Bolca Lagerstätten derives from a unique combination of age (near
shallow-water Eocene sequences deposited on the Lessini Shelf
the dawn of modern reef ecosystems), location (within an ancient
(sensu Bosellini 1989), a palaeogeographical feature of the
biodiversity hotspot) and taphonomy (exceptional preservation).
Southern Alps that was uplifted during the Alpine orogeny. Upon
Here we provide an overview of these sites, discussing history,
reaching the photic zone during the early Eocene, the Lessini Shelf
geology, fossils and significance.
acted as a centre of deposition of shallow-water carbonates (Doglioni
& Bosellini 1987; Bosellini 1989; Luciani 1989). Eruptive products
of extensive magmatic activity accumulated between the late
Geological context Paleocene and the middle Eocene (e.g. Macera et al. 2008).
Beginning in the early Eocene, these volcanic and volcanoclastic
Localities
deposits became intercalated with marls and limestones.
The village of Bolca lies on the eastern part of Monti Lessini not far Widespread regional faulting and cross-cutting volcanic units
from Verona, northern Italy. Monti Lessini represents a southern hinder stratigraphic studies of the Eocene succession of the Bolca
prolongation of the Southern Alps (Carminati et al. 2012). Several area. Consequently, the Eocene carbonates of the Lessini Shelf have
productive sites characterized by contrasting fossils are known from been collectively assigned to the ‘Calcari Nummulitici’, an informal
the Bolca region (Papazzoni et al. 2014; Fig. 1), and are collectively unit poorly constrained in terms of age and depositional setting.
known as ‘Monte Bolca’ in older literature although no such place Pesciara consists of a limestone block surrounded by volcanic
exists. We refer to this geographically compact collection of deposits (Fig. 1b). The outcrop is less than 20 m thick, covering an
environmentally and stratigraphically distinctive localities as the area of a few hundred square metres, and consists of a rhythmic
Bolca Lagerstätten, drawing distinctions between the various sites. alternation of finely laminated micritic limestone with fishes and
The most famous representative is Pesciara, which has been plants and grainstone bearing benthic fossils (Papazzoni &
exploited since the mid-16th century (Box 1) and yields exquisitely Trevisani 2006). Nearly all the fishes, soft-bodied invertebrates
preserved marine fishes, plants and soft-bodied invertebrates. It is and plants collected from Pesciara are from five bands of grey,
joined by Monte Postale, located nearby and also famous for marine laminated micritic limestone. The succession of Monte Postale
fishes and plants. These two localities are the source of most fossils consists of more than 130 m of grainstone that alternates with
from the Bolca area in museums, and have yielded over 100 000 massive coralgal limestone and laminated wackestone with fishes
exceptionally preserved fossils (e.g. Blot 1969; Box 1). This review and plants similar to those of Pesciara (e.g. Vescogni et al. 2016;
focuses on these two localities. Papazzoni et al. 2017). The laminated limestone varies in thickness
There are two additional nearby localities included among the depending on position in the succession. However, most layers are
Bolca Lagerstätten: Purga di Bolca and Vegroni. These freshwater about 1 m thick, very similar to those of Pesciara.
and brackish deposits are classically considered younger than Both the Pesciara and Monte Postale successions were deposited
Pesciara and Monte Postale, but their stratigraphic correlation with in the early late Ypresian. Calcareous nannofossils and larger
the more famous horizons is problematic (Papazzoni et al. 2014). benthic Foraminifera indicate that the Pesciara succession corre-
Purga di Bolca consists of lignites around a volcanic neck, and sponds to the upper part of that of Monte Postale. Monte Postale

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570 M. Friedman & G. Carnevale

Fig. 1. Location and geology of Pesciaria and Monte Postale, the principal Lagerstätten of Bolca. (a) Location of Bolca in Italy. (b) Topographic map of
region immediately NW of the village of Bolca. Colours indicate different geological units: yellow, Spilecco Limestone; green, laminated and bedded
limestone; blue, massive limestone; orange, volcanic rocks. Intensity of shading indicates either rock exposed in outcrop (dark) or inferred (light). Adapted
from Trevisani (2015).

spans the entire NP 13 and CNE 5 calcareous nannoplankton zones are rare, with palms represented only by fruits (complete palms are
(Papazzoni et al. 2017), corresponding to a large part of the Shallow found at Vegroni; Giusberti et al. 2014a). Leaves and leaflets,
Benthic Zone (SBZ) 11 in the time interval between c. 50.5 and especially of legumes, are the most common remains of dicots, but
48.96 Ma (see Agnini et al. 2014). This corresponds to the early twigs, fruits and flowers are also known. Specimens of whole plants
Eocene climatic optimum (EECO in the sense of Luciani et al. are known, but some of these are regarded as restorations or
2016). Pesciara has been assigned to the uppermost part of SBZ 11, composites (Wilde et al. 2014). In addition to these remains,
corresponding to the basal portion of NP 14 and CNE 6, between Pesciara also yields amber (Trevisani et al. 2005). As in the
48.96 and c. 48.5 Ma. macroflora, angiosperms dominate the microflora, with rare
gymnosperm pollen and fern spores (Kedves & Zsivin 1970).
In drawing comparisons with approximately coeval fossil floras
Floral and faunal composition
(e.g. lacustrine deposits of Messel), Wilde et al. (2014) noted lower
Study of different components of the Bolca fossil assemblages is diversity in the Pesciara–Monte Postale macroflora combined with a
uneven. The fishes of Pesciara and Monte Postale have been the distinctive composition in terms of both taxa and organs. They
subject of nearly continuous investigation for centuries. The so- attributed these features to a taphonomic filter arising from fluvial
called ‘minor fauna’, comprising non-fishes, of Pesciara and Monte transport to the ocean and subsequent shallow marine wave action,
Postale are poorly known by comparison (Carnevale et al. 2014; and also acknowledged the potentially important role of storms in
Giusberti et al. 2014b). Similarly, relatively little work has built transporting some plant remains.
upon foundational palaeobotanical studies from the 19th century
(Wilde et al. 2014).
Invertebrates
Plants Jellyfishes and corals
Marine plants and macroalgae Medusae from Pesciara comprise complete specimens preserving
delicate features (Giusberti et al. 2014b; Fig. 2d). Corals are a
Floating or fully marine monocots such as seagrasses are the most
common component of the benthic fauna, represented by broken
common elements of the Pesciara–Monte Postale flora (Wilde et al.
material in the coarse-grained carbonates intercalated between the
2014; Fig. 2a). Brown and red algae are represented by macrofossils
laminites and in the fossiliferous laminated wackestone of Monte
and biomarkers (Schwark et al. 2009).
Postale (Tang 2002; Marramà et al. 2016a; Fig. 1b and c).

Terrestrial plants
Ferns are absent, and gymnosperms are limited to long, needle-like Annelids
leaves. The terrestrial flora of Pesciara–Monte Postale is dominated Annelids from Bolca include four species of polychaete placed in
by angiosperms, specifically dicots (Fig. 2b). Terrestrial monocots two genera, and a single species of leech (Giusberti et al. 2014b).
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The Eocene Bolca Lagerstätten 571

Box 1. History of collecting at Bolca

The first documented report of a fossil from the Bolca Lagerstätten dates to the 16th century, and notes material belonging to the ambassador of the Holy Roman
Empire to the Venetian Republic (Mattioli 1550). Nearly a century later, the first illustration of a fossil from Bolca appeared in a catalogue of the collection of a
Veronese apothecary (Ceruti & Chiocco 1622; Fig. 2a).
Bolca fossils and their origin were extensively debated during the 18th century. It is also at this time that large collections were amassed by noblemen in Verona,
including Vincenzo Bozza, Ottavio Canossa and Giovanni Battista Gazola (Fig. 2b). By the end of 1791, Gazola’s own museum contained over a thousand well-
preserved fossil fishes, plus numerous plants and invertebrates. The abbot Giovanni Serafino Volta studied the Bozza collection and assigned most of the fishes to
modern tropical species in his lavishly illustrated catalogue (Volta 1796–1809).
The revolutionary armies of Napoleon confiscated about 600 fossils from the Gazola collection during the occupation of Verona in 1797. Subsequently transported
to Paris, these specimens were studied by de Blainville (1818) for an account in Nouveau Dictionnaire d’Histoire naturelle, and later by Louis Agassiz, who reviewed
Volta’s identifications (Agassiz 1835) and provided further descriptions in his monumental Recherches sur les Poissons fossiles (Agassiz 1833–1844). The first
detailed analysis of the fossil plants from Bolca was provided by Abramo Bartolomeo Massalongo in a series of monographic studies (e.g. Massalongo 1850, 1851,
1859) and further improved and expanded by Meschinelli & Squinabol (1892).
The most extensive collections of fossils from the Bolca Lagerstätten remain in Italy, particularly in Verona and Padua. Large collections outside Italy, such as
those in Paris, have interesting histories. Fossils at the Naturhistorisches Museum, Vienna, were presented by Massalongo to Emperor Franz Joseph I following an
assassination attempt in 1853; material at the Natural History Museum, London, derives largely from the purchased collections of William Willoughby Cole, 3rd Earl
of Enniskillen, and Sir Philip de Malpas Grey Egerton in the late 1800s; Bolca specimens at the Carnegie Museum of Natural History, Pittsburgh, were acquired in
1903 from the Belgian Baron de Bayet, executive secretary to King Leopold II. Fossils from Pesciara–Monte Postale are otherwise found in numerous smaller
collections throughout the world, often tracing their origins to early ‘cabinets of curiosities’.

Fig. 2. Historical aspects of palaeontology of the Bolca localities. (a) The first fossil fish to be figured from Bolca, the squirrelfish Berybolcensis
leptacanthus, MCSNV T.177 (top), with historical illustration from Ceruti & Chiocco (1622) (bottom). (b) One of two galleries dedicated to the
display of fossil fishes from Bolca in the museum of Giovanni Battista Gazola.

Arthropods Fishes
Pesciara and Monte Postale yield a diversity of terrestrial and marine Beginning with the first major accounts by Volta (1789, 1796–
arthropods (Fig. 3c and e). Insects include thysanuruans, odonatans, 1809), the fish fauna of Pesciara and Monte Postale has been the
trichopterans, coleopterans, orthopterans, heteropterans, hymenop- subject of numerous broad overviews (Blot 1980; Bannikov 2014;
terans and dipterans, with the last group being the most common Carnevale et al. 2014).
(Giusberti et al. 2014b). Other terrestrial arthropods include a
scorpion and possible pseudoscorpion. Marine crustaceans domin-
ate the minor fauna, and include three isopod genera, two Sharks and rays
stomatopod genera and 12 decapod genera (Giusberti et al. 2014b). Chondrichthyans are rare and little studied relative to the more
famous ray-finned fishes, comprising fewer than 20 species
Molluscs (Carnevale et al. 2014; Marramà et al. 2017a,b; Fig. 4a).
Nevertheless, the Pesciara–Monte Postale fauna is the most
Bivalves and gastropods are abundant in the coarse limestones
diverse assemblage of articulated chondrichthyans known from
between laminites, but are uncommon within the laminites
the Cenozoic. Batoids are represented by rhinobatids, narcinids,
themselves. However, six bivalve genera and some undescribed
platyrhinids, dasyatids, myliobatids and urolophids. Sharks include
forms, plus two gastropod genera, are reported from the laminites.
carchariniforms and a lamniform known only from teeth.
Rare cephalopods include decapodiform coleoids and a nautiloid
Chimaeroids are represented by a single isolated fin-spine
(Giusberti et al. 2014b).
belonging to the genus Ischyodus (Marramà et al. 2017a).
Extensive soft-tissue preservation in sharks visualized using
Lophophorates ultraviolet light (Fanti et al. 2016) indicates potential for
Lophophorates are limited to six terebratulid specimens and a single substantially more anatomical detail than available in historical
bryozoan from Pesciara (Giusberti et al. 2014b). accounts (Fig. 5).
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572 M. Friedman & G. Carnevale

Fig. 3. Plants and invertebrates from the Bolca Lagerstätten. (a) Seagrass similar to Phyllospadix or Posidonia, Museo Civico di Storia Naturale, Verona
(MCSNV) fB47. (b) Leaf generally assigned to the mallow Sterculia, MGP-PD (Museo di Geologia e Paleontologia, Università degli Studi di Padova) 20V.
(c) The spiny lobster Justinia desmaresti, MCSNV 23. (d) The medusa Simplicibranchia bolcensis, MCSNV m.B.2. (e) Hymenopteran insect, MCSNV
i.c.2NS. Scale bars represent 50 mm, with the exception of (e), where the scale bar represents 5 mm. Photographs courtesy of: plants, G. Roghi (Istituto di
Geoscienze e Georisorse, CNR, Padova); invertebrates, R. Zorzin (MCSNV).

Non-acanthomorph ray-finned fishes pleuronectiforms (flatfishes), siganids (rabbitfishes), scatophagids

Roughly 30 species constitute the non-acanthomorph ray-finned (scatties), priacanthids (bigeyes), sparids ( porgies), lutjanids
fishes of Pesciara–Monte Postale. These include late-surviving (snappers) and pomacentrids (damselfishes). Joining these taxa
pycnodonts, deep-bodied neopterygians abundant during the are groups associated with more open-water habitats, including
Mesozoic (Blot 1987; Fig. 4b). Other Mesozoic elements include beloniforms (halfbeaks and flying fishes), scombrids (tunas and
Platinx, the last known member of Crossognathiformes, an early mackerels), xiphioids (billfishes) and palaeorhynchids (an extinct
diverging teleost clade (Arratia & Tischlinger 2010; Sferco et al. billfish group), as well as several groups of uncertain relationships
2015). No fewer than nine families of eels represent Elopomorpha to modern lineages (e.g. exelliids).
(Blot 1978), and are joined by three osteoglossomorph species.
The latter are noteworthy as extant examples are freshwater, Terrestrial vertebrates
although several taxa are known from marine deposits in the
Paleogene (e.g. Bonde 2008). The most common fish specimens Reptiles
found in the Bolca Lagerstätten (Landini & Sorbini 1996, table 1), A turtle and two genera of boid snakes are known from Pesciara
clupeomorphs have only recently been studied in detail (e.g. (Carnevale et al. 2014). Reptiles are more common at Purga di
Marramà & Carnevale 2015a,b, 2016; Fig. 4c). Ostariophysans, a Bolca, and include crocodiles in need of taxonomic revision, a
dominant group of primarily freshwater fishes, include the snake and two genera of freshwater turtles (Giusberti et al. 2014a).
anatomically primitive Chanoides (Patterson 1984). No ‘prota-
canthopterygians’, stomiiforms (dragonfishes) or myctophiforms Birds
(lanternfishes) are known, and aulopiforms (lizardfishes and kin)
Birds are limited to isolated feathers from Pesciara. Whereabouts of
are represented by a single species (Marramà & Carnevale 2017;
skeletal remains reported in historical accounts are unknown
Fig. 4d).
(Carnevale et al. 2014).

Acanthomorphs
Taphonomy
Taxonomically, the ichthyofauna is dominated by spiny-rayed
teleosts, or acanthomorphs (Fig. 4e–l). Non-percomorphs from Fossils from Pesciara–Monte Postale, particularly the fishes, are
Bolca include lampridiforms, a zeiform and holocentroids usually found as part and counterpart with complete squamation,
(Carnevale et al. 2014; Davesne et al. 2017). Percomorphs traces of pigmentation and phosphatized soft tissues (e.g. Wilby &
make up the vast majority of fish species from the Bolca Briggs 1997; Fig. 5). Controlled excavations carried out in Pesciara
Lagerstätten, and include familiar inhabitants of modern reefs: and Monte Postale show that preservation and abundance of fossils
apogonids (cardinalfishes), acanthurids (surgeonfishes), caran- differ between the two sites as a consequence of contrasting
gids ( jacks), syngnathiforms ( pipefishes and allies), labrids depositional environments (Marramà et al. 2016a).
(wrasses), tetraodontiforms ( pufferfishes and allies), ephippids Exceptionally preserved fossils are abundant in the finely
(spadefishes), sphyraenids (barracudas), lophiiforms (anglerfishes), laminated micritic limestone of Pesciara, where they constitute the
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The Eocene Bolca Lagerstätten 573

Fig. 4. Fishes from the Bolca Lagerstätten. (a) The numbfish Titanonarke molini, Museo Civico di Storia Naturale, Verona (MCSNV) IG.VR.67290; (b)
the pycnodont Nursallia veronae, MCSNV II.D.173; (c) the round herring Trollichthys bolcensis, Natural History Museum, London (NHMUK) PV OR
37227; (d) the barracudina Holosteus esocinus, Museum national d’Histoire naturelle, Paris (MNHN) BOL 175; (e) the surgeonfish Eorandallius elegans,
MCSNV VIII.C.58; (f ) the moonfish Mene oblonga, Naturhistorisches Museum, Vienna (NHMW) 1853.XXVII.28; (g) the perch-like fish of unclear
affinities Ceratoichthys pinnatiformis, MCSNV T.950; (h) the cornetfish Urosphen dubia, NHMUK PV P 15638; (i) the batfish Eoplatax papilio, MCSNV
I.G. 24573; ( j) the mackerel Godsilia lanceolata, MCSNV T.89; (k) the monkfish Caruso brachysomus, MNHN BOL 42; (l) the triggerfish relative
Protobalistum imperiale, MCSNV T.21. Scale bars represent 50 mm.

vast majority of the collected specimens. The current taphonomic blooms might represent one of the main causes of death of marine
model for fishes invokes rapid accumulation of corpses on an anoxic organisms from Pesciara (Marramà et al. 2016a).
bottom, where a well-developed microbial biofilm delayed their Most fossils from Monte Postale are incomplete or strongly
decomposition, protected them from scavengers and bottom disarticulated (c. 90% of fish specimens) to the degree that they
currents, and promoted rapid mineralization. Tetany exhibited by cannot be identified at genus or species level. This incompleteness
numerous fish specimens (e.g. Fig. 4c) suggests that toxic algal and disarticulation, as well as the disruption of fins, S-shaped
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574 M. Friedman & G. Carnevale

in a silled depression with restricted bottom circulation, a short

distance from the coast at a depth of many tens of metres. According
to those researchers, coral reefs and seagrass beds were present in
close proximity. Based on an integrative approach using facies
analysis and foraminiferal palaeoecology, Papazzoni & Trevisani
(2006) proposed deposition in a subtropical lagoon close to an
emerged area and characterized by seasonal changes of water
circulation that modulated oxygen concentrations near the seafloor.
What is agreed upon by all the recent studies is that sediments of
Pesciara reflect a peri-reefal system subject to the ecological
influence of both coastal environments and the open sea, and that
the deposition of the fossiliferous micritic limestone took place in a
low-energy basin characterized by permanent bottom dysoxia or
anoxia (Marramà et al. 2016a). Sometimes conspicuous primary
production was dominated by diatoms (Schwark et al. 2009), which
constituted the base of the trophic chain of the Pesciara assemblage
and sustained large shoals of the sardine Bolcaichthys catopygop-
terus (Marramà & Carnevale 2015a).
Roughly half of fossil specimens from the laminated wackestone
of Monte Postale are remains of algae and plants, followed by fishes
(about 30%) showing different degrees of completeness, and
abundant invertebrates (corals, molluscs and crustaceans). Fishes
primarily consist of small epibenthic species and rarer large taxa
(Marramà et al. 2016a). The laminated wackestone of Monte Postale
originated in a lagoon surrounded by coralgal buildups made of
corals, calcareous algae and encrusting foraminifera (Vescogni et al.
2016). The peri-reefal areas were densely vegetated by seagrass beds
and mangroves, the latter dominated by the mangrove palm Nypa
(Marramà et al. 2016a).

Evolutionary and palaeoecological significance

The Bolca Lagerstätten provide critical snapshots of reef-associated
marine ecosystems and adjacent terrestrial areas in the early
Cenozoic. As such, they are a unique line of evidence bearing on
the origin and early history of modern reefs, which represent major
hotspots of marine biodiversity. As a result of previous research, the
importance of the Bolca localities is clearest for fishes and derives
Fig. 5. Examples of exceptional preservation of fossils from the Bolca from three main features apart from excellent preservation: early
Lagerstätten. (a) Preservation of pigmentation ( p), soft tissues and gut Eocene age, unique environmental setting and palaeogeographical
contents in the triakid shark Galeorhinus cuvieri, Museo Geologico location. With respect to age, the principal Bolca sites postdate the
Giovanni Capellini, Bologna (MGGC) 1976; specimen shown in right Cretaceous–Paleogene (K–Pg) mass extinction by roughly 15 myr,
lateral view and illuminated in ultraviolet light. Dashed-line rectangle providing a glimpse into a mature ecosystem following an interval
indicates area shown in (b). (b) Close-up view of abdominal cavity of of evolutionary recovery (see Erwin 2001; Box 2, Fig. 6). With
specimen in (a), including spiral valve of intestine (sv) and skeleton of the respect to environmental setting, Pesciara–Monte Postale show
barracuda Sphyraena bolcensis representing gut contents (gc). Images substantially higher species richness of fishes than, and contrasting
courtesy of T. Miyashita (University of Alberta). (c) Pigmentation patterns
taxonomic composition to, other Ypresian deposits that can be
in the eel Paranguilla tigrina, Museo di Geologia e Paleontologia,
considered Lagerstätten in their own right (Danata Formation,
Università degli Studi di Padova (MGP-PD) 26288; specimen shown in
left lateral view. Image courtesy of S. Castelli and L. Giusberti (MGP- Turkmenistan; Fur Formation, Denmark; London Clay Formation,
PD). Scale bars: (a) 75 mm; (b) 25 mm; (c) 20 mm. UK; Friedman et al. 2016); it is possible that this is due to close
association with reefs. With respect to palaeobiogeography, the
Bolca sites are ideally positioned for capturing a high-diversity
curving of the vertebral column and the unidirectional dispersion of marine assemblage of Paleogene age. High species richness of
scales around the body are indicative of episodic disturbance and benthic foraminifera, corals and molluscs in the late Eocene of the
mixing of the bottom. These occasional events resulted in the western Tethys, the area including Bolca, suggests that this region
development of normal aerobic conditions on the bottom that was an ancient biodiversity hotspot, comparable with today’s Indo-
eventually allowed the settling of a moderately diverse benthic Pacific (Renema et al. 2008; Fig. 7a).
fauna, revealed by relatively abundant bioturbation tracks and Modern coralgal reef ecosystems trace their origins to the early
crustacean and mollusc remains (Marramà et al. 2016a). Paleogene (Kiessling 2009; Bellwood et al. 2017). Temporally, the
Bolca Lagerstätten fall near the end of a second wave of reef
Palaeoenvironment invasions by modern teleosts as inferred on the basis of comparative
phylogenetic data (Price et al. 2014), and their fauna is often hailed
The differences in taphonomy and of the fish assemblages as the earliest modern reef-fish assemblage (Bellwood 1996;
themselves suggest different palaeoenvironmental settings for the Bellwood & Wainwright 2002). Such comparisons are not without
two principal Bolca localities. Landini & Sorbini (1996) hypothe- merit, but there are differences from living reef faunas (Fig. 7b–d).
sized that deposition of the laminated limestone of Pesciara occurred Some major groups of reef-associated fishes such as butterflyfishes
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The Eocene Bolca Lagerstätten 575

Box 2. Bolca and the evolutionary radiation of acanthomorphs

Acanthomorphs, or the spiny-rayed teleosts, include nearly one in three living vertebrate species and the majority of fish species. Their diversity in form is
commensurate with their diversity in kind, with modern acanthomorphs assuming morphologies as divergent as those of flounders, seahorses, anglerfishes, tunas and
mudskippers. The majority of acanthomorphs in turn belong to a group known as percomorphs, a species-rich radiation that has long vexed systematists as the
unresolved ‘bush at the top’ of the teleost tree of life. The first acanthomorphs appear in the fossil record at the beginning of the Late Cretaceous, around 100 myr ago,
represented by deeply branching lineages such as lampridiforms and beryciforms (Davesne et al. 2014, 2016; Fig. 6). The oldest definitive percomorphs make their
debut tens of millions of years later, near the end of the Late Cretaceous, and belong to early diverging groups such as syngnathiforms, ophidiiforms and possibly
batrachoidiformes (Carnevale & Johnson 2015). Although some basal phylogenetic splits had clearly taken place by the Late Cretaceous, acanthomorphs remained
comparatively minor components of marine faunas of this age, rarely making up more than a third of all fish species at any given fossil locality (Patterson 1993a;
Friedman et al. 2016). Highly selective extinction among marine fishes occurred at the end of the Cretaceous (Friedman 2009). Faunas of the earliest Cenozoic remain
poorly known, but largely include acanthomorph groups already known from the Late Cretaceous (Alvarado-Ortega et al. 2015; Adolfssen et al. 2017). A series of
deposits from near or shortly after the Paleocene–Eocene Thermal Maximum in Europe and western Asia provide the first evidence of acanthomorph-dominated
faunas with highly diversified percomorph lineages (Patterson 1993a; Friedman et al. 2016; Bannikov et al. 2017). For the most part, these deposits represent open-
water settings, with abundant representatives of pelagic groups but showing only modest diversity of nearshore or reef-associated lineages. Pesciara–Monte Postale
postdate the earliest of these assemblages by roughly 6 myr, and further mark the shift to percomorph-dominated marine fish faunas. Significantly, the Bolca
localities capture the first (and in a few cases, only) body fossil occurrences of many reef-fish clades (Fig. 6). Increased taxonomic dominance of acanthomorphs in
the early Paleogene is associated with a substantial increase in morphological diversity as measured by overall body shape (Friedman 2010), although there is some
evidence for an earlier onset of cranial diversification (Sallan & Friedman 2012). Pesciara and Monte Postale are unquestionably exceptional, but the pattern of
increased anatomical diversity among acanthomorphs in the early Paleogene appears robust to the exclusion of these sites from quantitative analyses (Friedman
2010).

Fig. 6. Time-calibrated molecular phylogeny of extant acanthomorph teleosts, adapted from Near et al. (2013). Percomorph acanthomorphs
highlighted in orange. Concentric rings represent Cretaceous–Paleogene (K–Pg) boundary (dashed line) and the age of the principal fish-bearing Bolca
localities of Pesciara and Monte Postale (continuous black line). Filled circles indicate extant family-level clades known from Bolca. A black filled
circle indicates that material from these sites represents the first body-fossil appearance for the group in the geological record.

and parrotfishes are not found at Pesciara–Monte Postale, and only (Bellwood 1996; Bellwood et al. 2017; Fig. 7c). There are
appear later in the Cenozoic (Bellwood & Schultz 1988; Carnevale varying degrees of overlap between fossils of the Bolca
2006). Associated with the lack of some modern groups are Lagerstätten and their modern reef-dwelling relatives, as gauged
remarkable examples of evolutionary convergence between Eocene by geometric morphometric quantification of body shape (Bellwood
and extant reef fishes (Tyler & Bannikov 2005). Contrasts in terms of et al. 2014a; Marramà et al. 2016b), as well as differences in
diversity of genus-level lineages within families, as well as relative ecomorphologically relevant traits such as eye size (Goatley et al.
abundance of individuals within families, are also apparent 2010), mandibular mechanics (Bellwood 2003; Bellwood & Hoey
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576 M. Friedman & G. Carnevale

Fig. 7. The fossil fishes of the Bolca Lagerstätten represent an early coral reef assemblage. In panels comparing multiple localities, Bolca is represented by
a star. (a) Patterns of biodiversity in Tethys during the Eocene as inferred based on benthic Foraminifera. The area including Bolca represents a ‘hotspot’
enclosed by dashed line. Adapted from Renema et al. (2008). (b) Principal components ordination of modern reef-fish assemblages and the Pesciara–Monte
Postale ichthyofauna, based on taxonomic composition. Adapted from Bellwood (1997). (c) Differences between genus-level diversity within major extant
reef-fish families in a modern rocky reef in the eastern Pacific (top) and Pesciara–Monte Postale (bottom). Other modern reef ecosystems show similar
contrasts to these Eocene sites. Adapted from Bellwood et al. (2017). (d) Morphological disparity (multivariate variance) of modern reef-fish faunas and
that of Pesciara–Monte Postale, assessed from a geometric morphometric analysis of body shape and fin position. Adapted from Marramà et al. (2016b).

2004), jaw protrusion (Bellwood et al. 2015) and dental morphology To a first order of approximation, the Pesciara–Monte Postale
(Bellwood et al. 2014b). Although commonly compared with Indo- fauna appears modern and there is much truth to the quip of
Pacific assemblages on the basis of groups that are endemic to this Patterson (1993a, p. 53) that subsequent evolution among marine
modern biodiversity hotspot, the Pesciara–Monte Postale fish fauna fishes ‘was mere tinkering’ (Box 2). However, there are peculiar
is not strictly comparable with those of either the Atlantic or Indo- extinct phenotypes among otherwise familiar bodyplans. These
Pacific (Bellwood 1997; Fig. 7b). Taken together with other include: early relatives of flatfishes indicating gradual, rather than
localities, the Bolca sites provide a picture of a widespread saltational, evolution of asymmetry (Friedman 2008, 2012), although
Paleogene marine fish fauna. At higher taxonomic levels (i.e. that stepwise change might have occurred over a geologically short
family), it is apparent that the Indo-Pacific region has maintained interval (Harrington et al. 2016); early relatives of gymnodont
much of this diversity, whereas Atlantic reef assemblages have been tetraodontiforms (Tyler & Santini 2002) that provide clues to the
winnowed by regional extinction (Bellwood 1997). evolutionary origin of the beak-like dentition further elaborated by
Fossils from the Bolca Lagerstätten also deliver important living members of the group (Fraser et al. 2012); the peculiar
evidence relating to individual fish clades, especially in terms of †Bajaichthys, a zeiform (John Dory) that shows an unprecedented
constraints on times of evolutionary divergence and detailed ribbon-like morphology in a group otherwise characterized by deep
character data bearing on patterns of phenotypic evolution. The bodies (Davesne et al. 2017); the extinct syngnathiform group
oldest occurrences of many teleost families are from Pesciara– †Aulorhamphidae, which combines the elongated snouts so charac-
Monte Postale (Patterson 1993a,b; Fig. 6), and fishes from these teristic of the order with generalized postcrania (Tyler 2004).
localities serve as important temporal constraints for estimating
evolutionary timescales under a variety of approaches (e.g. Near
Summary
et al. 2012, 2013; Chen et al. 2014; Arcila et al. 2015; Close et al.
2016; Bannikov et al. 2017). The impact of Pesciara–Monte Postale The Bolca Lagerstätten provide a unique window into a series of
on raw counts of teleost familial diversity through the Cenozoic is fully marine to terrestrial environments in the early Eocene
striking, and suggestive of an extreme Lagerstätten effect (Patterson of Tethys. Their remarkable diversity reflects a fortuitous
1994). Such obvious bias aside, the Bolca Lagerstätten and other combination of taphonomic factors combined with a reef setting
fossil deposits of similar age all record a major shift in marine fish in a region interpreted as an ancient biodiversity hotspot. The most
assemblages between the Late Cretaceous and Paleogene, defined celebrated sites of the Bolca Lagerstätten are Pesciara and Monte
most conspicuously by the increased taxonomic dominance of Postale, both of which yield exquisitely preserved fish fossils
acanthomorphs (spiny-rayed fishes) in general and percomorphs totalling well over 200 species. The fossil fishes from these sites are
( perch-like fishes) in particular (Box 2). particularly significant in documenting the rise of acanthomorphs,
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The Eocene Bolca Lagerstätten 577

Box 3. Outstanding questions

The past few years have seen substantial improvements in our understanding of the geological and environmental context of the Pesciara and Monte Postale sites (e.g.
Marramà et al. 2016a), coupled with increasingly sophisticated quantitative analysis of their fish faunas (e.g. Bellwood et al. 2014a; Marramà et al. 2016b). However,
a series of outstanding challenges must still be addressed. First, clear stratigraphic correlation between freshwater to brackish Vegroni and Purga di Bolca localities
and classic marine sites of Bolca remain elusive, partly owing to the complex post-depositional geological history of the region. Clarity on the age of these nearshore
to terrestrial deposits will provide insight into the degree to which the Bolca sites collectively represent a geological snapshot, or are instead diachronous. Second,
many components of the extended Bolca assemblage are in dire need of taxonomic revision. Nowhere is this more conspicuous than among plants, known from each
of the Bolca localities but subject to remarkably little targeted investigation over the past century. Third, although the fishes are without question the best known
taxonomic component of the Bolca assemblage, it could be argued that they remain understudied relative to the amount of available material. This issue is more acute
for some groups (e.g. chondrichthyans), but it is clear that more information is preserved in material studied in the past, from patterns of pigmentation, soft-tissue
preservation, and fine osteological details accessible through transfer preparation. New anatomical data for old fossils might help better constrain their phylogenetic
positions, a prerequisite for higher-level analyses addressing macroecological and macroevolutionary questions surrounding the origins of modern reef-fish
assemblages and specific clades (Box 2).

the dominant group of marine teleosts, as well as providing critical Bellwood, D.R. & Hoey, A.S. 2004. Feeding in Mesozoic fishes: a functional
clues to the early assembly of modern tropical reef ecosystems. perspective. In: Arratia, G. & Tintori, A. (eds) Mesozoic Fishes 3 –
Systematics, Paleoenvironments and Biodiversity. Pfeil, Munich,
Although they are often overshadowed by the fishes, a wide range of 639–649.
marine invertebrates, aquatic plants and algae, terrestrial plants and Bellwood, D.R. & Schultz, O. 1988. A review of the fossil record of the
insects are also present at Pesciara and Monte Postale. The marginal parrotfishes (Labroidei: Scaridae) with a description of a new Calotomus
species from the Middle Miocene (Badenian) of Austria. Annalen des
to terrestrial sites near Bolca remain little examined, and their Naturhistorischen Museums in Wien, Serie A, 92, 55–71.
relationship to the ‘classic’ localities remains unclear (Box 3). Bellwood, D.R. & Wainwright, P.C. 2002. The history and biogeography of
fishes on coral reefs. In: Sale, P.F. (ed.) Coral Reef Fishes. Elsevier,
Amsterdam, 5–32.
Acknowledgements We thank P. Donoghue (University of Bristol) for Bellwood, D.R., Goatley, C.H.R., Brandl, S.J. & Bellwood, O. 2014a. Fifty
his invitation to write this review, and our many colleagues studying the fossils million years of herbivory on coral reefs: fossils, fish and functional
and geology of the Bolca sites for their helpful input over the years. We especially innovations. Proceedings of the Royal Society of London, Series B, 281,
thank the Museo Civico di Storia Naturale, Verona, the Museo di Geologia 20133046.
e Paleontologia, Università degli Studi di Padova, Padua, Muséum national Bellwood, D.R., Hoey, A.S., Bellwood, O. & Goatley, C.H. 2014b. Evolution of
d’Histoire naturelle, Paris, the Natural History Museum, London and long-toothed fishes and the changing nature of fish–benthos interactions on
Naturhistorisches Museum, Vienna for allowing us to photograph fossil fishes coral reefs. Nature Communications, 5, 3144.
in their collections. C. Abraczinskas (University of Michigan Museum of Bellwood, D.R., Goatley, C.H., Bellwood, O., Delbarre, D.J. & Friedman, M.
Paleontology) produced or improved the figures. A. Capobianco (University of 2015. The rise of jaw protrusion in spiny-rayed fishes closes the gap on elusive
Michigan), D. Harper (University of Durham) and an anonymous referee prey. Current Biology, 25, 2696–2700.
provided helpful comments on an earlier version of this paper. Bellwood, D.R., Goatley, C.H. & Bellwood, O. 2017. The evolution of fishes and
corals on reefs: form, function and interdependence. Biological Reviews, 92,
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