The systematic composition of the Eocene

Studi e ricerche sui giacimenti terziari di Bolca, XV - Miscellanea paleontologica, 12, 2014: 23 - 34
The systematic composition of the Eocene actinopterygian fish fauna
from Monte Bolca, northern Italy, as known to date
ALEXANDRE F. BANNIKOV
(Borisyak Paleontological Institute of the Russian Academy of Sciences, Moscow)
Abstract
The most recent list of fishes from the especially rich Cenozoic Monte Bolca locality (northern Italy) was that of Blot (1980). Since that
time, the systematic composition of this late Ypresian fish fauna has been much revised and substantially expanded by the contributions
of various scientists in numerous publications, and to date these actinopterygian fishes comprise 19 orders, 91 families, 192 genera (32 of
which are of uncertain family) and at least 220 species. Among these, the Acanthomorpha comprise 11 orders, 78 families, 163 genera
and at least 190 species.
Key words: Actinopterygii, Acanthomorpha, taxonomy, Lower Eocene, northern Italy, Monte Bolca locality.
Riassunto
L’ultima lista di pesci fossili del ricchissimo giacimento cenozoico di Bolca (Italia settentrionale) fu pubblicata da Blot (1980).
Successivamente la composizione sistematica dell’ittiofauna dell’Ypresiano superiore di Bolca fu revisionata e grandemente completata dai contributi di numerosi autori in svariate pubblicazioni e, al momento attuale, consiste di 19 ordini, 91 famiglie, 192
generi (32 dei quali rappresentanti famiglie di posizione incerta) e almeno 220 specie di attinopterigi; fra questi 11 ordini, 78
famiglie, 163 generi e almeno 190 specie appartengono agli Acanthomorpha.
Parole chiave: Actinopterygii, Acanthomorpha, tassonomia, Eocene Inferiore, Italia settentrionale, Bolca.
The fish fauna of the Monte Bolca locality in northern Italy is the most diverse of all of the Cenozoic ichthyofaunas; it is the most comprehensive shallow-water Tethian fish fauna known for the first half of the
Eocene. The fishes of Monte Bolca have been known
since the XVIth century, with the specimens collected from two sites: the Pesciara, where the fossils are extracted from caves, and Monte Postale, where these are
sampled directly from the outcrop. Medizza (1975) dated the Monte Bolca locality as the uppermost Lower to
the lowermost Middle Eocene (Discoaster sublodoensis
Zone), whereas Papazzoni and Trevisani (2006) determined a late Ypreseian (Alveolina dainellii Zone) age for
the Pesciara (about 50 mya).
Unlike most of the other known assemblages of marine fossil fishes, which usually reflect relatively well only
the composition of the pelagic component of the fauna,
in the Monte Bolca assemblage bottom dwellers are also
relatively well represented. The bottom fishes of Monte
Bolca are representatives of the extant families Congridae, Ophichthyidae, Ophidiidae, Lophiidae, Brachionichthyidae, Antennariidae, and Ogcocephalidae. Other
bottom dwellers are the Eocene genera of lampridiforms (Bajaichthys Sorbini, 1983) and pleuronectiforms
(Eobothus Eastman, 1914, Amphistium Agassiz, 1844 and
Heteronectes Friedman, 2008), as well as the fossil family
Eocottidae, whose representatives in the Eocene fauna
of Monte Bolca are the ecological analogues of the Recent gobies (Gobiidae) and sculpins (Cottidae) (Bannikov, 2004a). The Monte Bolca locality represents the
most ancient, richest, and best known assemblage of
coral reef fishes of the modern type (Bellwood, 1996),
with perciforms predominating. Unlike Recent coral reef ichthyofaunas, in which pomacentrids, apogonids, gobiids, labrids and chaetodontids usually predominate, in the Eocene coral reef associated fauna of
Monte Bolca it is holocentrids that are most commonly collected, and acanthurids are extremely diverse taxonomically (14 genera). At Monte Bolca, the family Apogonidae and suborder Labroidei each are represented
by only five or six monotypic genera and the Pomacentridae are exceptionally rare. The unquestionable early Eocene Gobiidae and Chaetodontidae are unknown
at Monte Bolca to date. The pycnodontid genera Pycnodus Agassiz, Palaeobalistum Blainville, Nursallia Blot
and Abdobalistum Poyato-Ariza et Wenz (the last indication of the Pycnodontiformes in the paleontological
record) are relicts within the Monte Bolca assemblage
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ALEXANDRE F. BANNIKOV
of the Cretaceous coral fish faunas. The ichthyofauna
of Monte Bolca differs from that at the geographically nearby Monte Solane locality (Zorzin et al., 2011),
which is of similar age to Monte Bolca, by having no
mesopelagic fishes and relatively few unquestionably
epipelagic fishes (such as euzaphlegids, scombrids and
palaeorhynchids). Extant paralepidids are mesopelagic
fishes, but the species of the fossil genus Holosteus Ag.
are known from only relatively shallow water habits.
Papazzoni and Trevisani (2006) reconstructed the environment of the Pesciara Lagerstätte (the main site of the
Bolca locality) as a restricted shallow “basin” in which
the interchange of water was very low, affecting the oxygen content. In this basin, sedimentation was mainly
micritic, possibly with occasional stromatolite-like bacterial mats. The basin was affected by sea-level oscillations and coarser grained storm-induced deposition.
The first monograph with descriptions of Bolca fishes was published toward the end of the XVIIIth century (Volta, 1796). Volta described about 100 species,
most of which were erroneously identified as conspecific with extant taxa, and only two genera, Monopteros and Blochius, and 15 species were described as new.
Subsequently, many of Volta’s determinations were improved by Blainville (1818). Most of the specimens described by Volta as being in Italy were soon thereafter
removed to Paris by order of Napoléon Bonaparte, and
these were re-described there by Agassiz (1833-1844).
Agassiz recognized the only species of the genus Blochius described by Volta, B. longirostris, as having been adequately established; and described all of the other 14
species as new or with the authorship credited to Blainville. The list of fishes from Monte Bolca was increased
by the descriptions of new taxa in the XIXth century
(contributions of Heckel, de Zigno, Bassani, etc.) and
in the early XXth century (Woodward, 1901; Eastman,
1904, 1905), and consisted of at least 135 nominal species
of bony fishes (Leriche, 1906). It is pertinent to note
here that among the old Bolca materials some skeletons
were composed by the preparators from the parts of different species of fishes, often of different genera. These
skeletons are falsified in a scientific sense and it is often difficult to recognize them. For example, the specimen described by Volta (1796: 193, pl. XLVIII, fig. 1)
as Ophicephalus striatus and regarded usually as a synonym of the scombrid Auxides propterygius (e.g., Woodward, 1901: 464), and presently housed in the Museum
of Le Mans, is composed of the skeletons of at least two
different species; most of the skeleton belongs to Callipteryx, whereas its anterior portion is of Sphyraena.
In the second half of the XXth century, primarily
due to the contributions of J. Blot and L. Sorbini, a
goodly number of the names of Bolca species were synonymized and many new genera and species were described. The list of fishes from Monte Bolca published
by Blot (1980) includes 72 families, 117 genera (plus a
number of listed but as yet undescribed genera) and
208 species (including as yet undescribed species) of
bony fishes. Subsequently, the systematic composition
of the Monte Bolca fish fauna was revised and greatly
expanded by the contributions of many scientists (e.g.,
Sorbini, 1983; Bannikov, 1991, 1999, 2000, 2004a, b,
2005, 2006, 2008a-f; Bannikov and Sorbini, 1991; Blot
and Tyler, 1991; Bannikov and Tyler, 1999; Bellwood,
1999; Tyler, 1999; Tyler and Bannikov, 2002, 2005, 2011;
Monsch, 2006; Bannikov and Carnevale, 2009, 2010,
2011; Carnevale and Pietsch, 2009, 2011; Bannikov and
Zorzin, 2011, 2013; Pietsch and Carnevale, 2011; etc.).
The extant perciform families Serranidae, Moronidae, Percidae, Terapontidae, Haemulidae (=Pristipomidae), Chaetodontidae, Enoplosidae, Mullidae, Trachinidae and Blenniidae indicated by Blot (1980) in
his systematic catalogue have to be excluded from the
Bolca ichthyofauna, as follows. It was shown that the
specimen of the labroid Sorbinia caudopunctata Bellwood housed in the Museo Civico di Storia Naturale
di Verona (MCSNV IG24561) was erroneously classified by Blot with the Mullidae (Bannikov, 2008b); the
“serranid” Serranus rugosus Heckel is actually a synonym of Veranichthys ventralis (Ag.) (Lutjanidae), whereas the “haemulid” Pristipoma furcatum Ag. is a synonym of Sparnodus elongatus Ag. (Sparidae) (Bannikov,
2006); the “moronid” Labrax schisurus Ag. is the type
of a new genus of Percoidei inc. sed., Veronabrax (Bannikov, 2008a); the “percid” Dules temnopterus Ag. and
the “terapontid” Pelates quindecimalis Ag. also belong
to Percoidei inc. sed., to the genera Jimtylerius and Parapelates, respectively (Bannikov and Carnevale, 2007;
Bannikov, 2008c). The genus Callipteryx should be
transferred from the Trachinidae into the family Callipterygidae Jordan, and the genus Oncolepis from the
Blenniidae into the Acanthopterygii inc. sed. The species Enoplosus pygopterus Ag. was transferred from the
Enoplosidae into the Apogonidae, as the genus Eosphaeramia Sorbini, 1983; and Eosphaeramia margaritae Sorbini, 1983 is its synonym (Bannikov, 2008d). A
review of the fishes from Monte Bolca that previously
had been referred to the Chaetodontidae revealed (Bannikov, 2004b) that none of them is actually related to
the butterflyfishes: the genus Acanthopygaeus Leriche,
1906 should be regarded as a member of the Siganidae (see also Bannikov et al., 2010), the genera Pygaeus
Agassiz, 1838 and Malacopygaeus Leriche, 1906 as Percoidei inc. sed., and the genus Parapygaeus Pellégrin,
The systematic composition of the Eocene actinopterygian fish fauna from Monte Bolca, northern Italy
1907 as Perciformes inc. sed. Blot (1980) indicated four
specimens in Verona as the types for new genera and
species of chaetodontids to be described by him, but his
untimely death prevented the fulfillment of that plan.
Close examination of those specimens at the MCSNV
indicates that none of them actually represents a chaetodontid (Bannikov, 2004b: 57).
A number of new fossil genera were established by
various authors during the last 30 years, some of them
in order to replace the names of extant taxa in the Bolca
fish fauna. Moreover, the systematic position of many
taxa was corrected, and some old generic and species
names were regarded as synonyms. And the additional taxa of fishes described during that period included representatives of families that had not previously
been known from Bolca, as follows: the extant families Chlopsidae, Veliferidae, Antennariidae, Gerreidae,
Apolectidae, Leiognathidae, Lutjanidae, Caproidae,
Siganidae, Centrolophidae and extinct families Pachyrhizodontidae, Euzaphlegidae and Palaeorhynchidae;
and the new fossil families Foreyichthyidae, Bajaichthyidae, Rhamphognathidae, Mesogasteridae, Aulorhamphidae, Quasimullidae, Eocottidae, Robertanniidae, Tortonesidae, Sorbinipercidae, Zorzinichthyidae,
Acanthonemidae, Massalongiidae, and Bolcabalistidae.
As of now, the fish fauna of Monte Bolca includes
the following taxa of actinopterygian fishes (the systematic arrangement follows Nelson, 1994, 2006; with
some emendations):
Division HALECOSTOMI
Order Pycnodontiformes
Family Pycnodontidae
Genus Pycnodus Agassiz, 1833
Pycnodus apodus (Volta, 1796)
Genus Nursallia Blot, 1987
Nursallia veronae Blot, 1987
Genus Abdobalistum Poyato-Ariza et Wenz, 2002
Abdobalistum thyrsus Poyato-Ariza et Wenz, 2002
Genus Palaeobalistum Blainville, 1818
Palaeobalistum orbiculatum Blainville, 1818
Division TELEOSTEI
TELEOSTEI inc. sed.
Order Crossognathiformes
Family Pachyrhizodontidae
Genus Platinx Agassiz, 1835
Platinx macropterus (Blainville, 1818)
Subdivision OSTEOGLOSSOMORPHA
Order Osteoglossiformes
Family Foreyichthyidae
Genus Foreyichthys Taverne, 1979
Foreyichthys bolcensis Taverne, 1979
Family Arapaimidae
Genus Thrissopterus Heckel, 1856
Thrissopterus catullii Heckel, 1856
Osteoglossiformes inc. sed.
Genus Monopteros Volta, 1796
Monopteros gigas Volta, 1796
Subdivision ELOPOMORPHA
Order Anguilliformes
Family Anguilloididae
Genus Anguilloides Cadrobbi, 1962
Anguilloides branchiostegalis (Eastman, 1905)
Genus Veronanguilla Blot, 1978
Veronanguilla ruffoi Blot, 1978
Family Milananguillidae
Genus Milananguilla Blot, 1978
Milananguilla lehmani Blot, 1978
Family Anguillidae
Genus Eoanguilla Blot, 1978
Eoanguilla leptoptera (Agassiz, 1835)
Family Paranguillidae
Genus Paranguilla Bleeker, 1864
Paranguilla tigrina (Agassiz, 1839)
Genus Dalpiaziella Cadrobbi, 1962
Dalpiaziella brevicauda Cadrobbi, 1962
Family Congridae
Genus Voltaconger Blot, 1978
Voltaconger latispinus (Agassiz, 1839)
Genus Bolcyrus Blot, 1978
Bolcyrus formosissimus (Eastman, 1905)
Bolcyrus bajai Blot, 1978
Genus Paracongroides Blot, 1978
Paracongroides heckeli Blot, 1978
Family Chlopsidae
Genus Whitapodus Blot, 1980
Whitapodus breviculus (Agassiz, 1835)
Family Proteomyridae
Genus Proteomyrus Cadrobbi, 1962
Proteomyrus ventralis (Agassiz, 1839)
Family Ophichthyidae
Genus Goslinophis Blot, 1980
Goslinophis acuticaudus (Agassiz, 1835)
Family Patavichthyidae
Genus Patavichthys Blot, 1980
Patavichthys bolcensis (Bassani, 1897)
Anguilliformes inc. sed.
Genus Bolcanguilla Blot, 1980
Bolcanguilla brachycephala Blot, 1980
Genus Gazolapodus Blot, 1980
Gazolapodus homopterus Blot, 1980
Subdivision CLUPEOMORPHA
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ALEXANDRE F. BANNIKOV
Order Clupeiformes
Family Clupeidae
Genus “Clupea”
“Clupea” catopygoptera Woodward, 1901
Genus Trollichthys Marramà et Carnevale, MS
Trollichthys bolcensis Marramà et Carnevale, MS
Nomina dubia:
“Clupea” cephalus Lioy, 1866
“Clupea” chrysosoma Lioy, 1866
“Clupea” denticiformis Lioy, 1866
“Clupea” engrauliformis Lioy, 1866
“Clupea” microcephala Lioy, 1866
“Clupea” minutissima Lioy, 1866
“Clupea” ophthalmica Lioy, 1866
“Clupea” polyachanthina Lioy, 1866
Ptericephalina elongata Lioy, 1866
Ptericephalina macrogastrina Lioy, 1866
Uropterina platyrachis Lioy, 1866
Subdivision EUTELEOSTEI
Superorder Ostariophysi
Series Anotophysi
Anotophysi inc. sed.
Genus Coelogaster Eastman, 1905
Coelogaster leptostea (Eastman, 1905)
Genus “Chanos”
“Chanos” forcipatus (Heckel, 1854)
Series Otophysi
Otophysi inc. sed.
Family Chanoididae
Genus Chanoides Woodward, 1901
Chanoides macropoma (Agassiz, 1844)
Superorder Cyclosquamata
Order Aulopiformes
Family Paralepididae
Genus Holosteus Agassiz, 1839
Holosteus esocinus Agassiz, 1839
Superorder Lampridiomorpha
Order Lampridiformes
Family Veliferidae
Genus Veronavelifer Bannikov, 1990
Veronavelifer sorbinii Bannikov, 1990
Genus Velifer Temminck et Schlegel, 1850
Velifer carosii Carnevale et Bannikov, MS
Family Bajaichthyidae
Genus Bajaichthys Sorbini, 1983
Bajaichthys elegans Sorbini, 1983
Lampridiformes inc. sed.
Genus “Pegasus”
“Pegasus” volans Volta, 1796
Superorder Paracanthopterygii
Order Ophidiiformes
Family Ophidiidae
Genus “Ophidium”
“Ophidium” voltianum Massalongo, 1859
Order Lophiiformes
Suborder Lophioidei
Family Lophiidae
Genus Sharfia Pietsch et Carnevale, 2011
Sharfia mirabilis Pietsch et Carnevale, 2011
Genus Caruso Carnevale et Pietsch, 2012
Caruso brachysomus (Agassiz, 1839)
Suborder Antennarioidei
Family Antennariidae
Genus Eophryne Carnevale et Pietsch, 2009
Eophryne barbutii Carnevale et Pietsch, 2009
Family Brachionichthyidae
Genus Histionotophorus Eastman, 1904
Histionotophorus bassanii (de Zigno, 1887)
Genus Orrichthys Carnevale et Pietsch, 2010
Orrichthys longimanus Carnevale et Pietsch, 2010
Suborder Ogcocephalioidei
Family Ogcocephalidae
Genus Tarkus Carnevale et Pietsch, 2011
Tarkus squirei Carnevale et Pietsch, 2011
Superorder Acanthopterygii
Series Atherinomorpha
Order Atheriniformes
Family Atherinidae
Genus Atherina Linnaeus, 1758
Atherina (?) macrocephala Woodward, 1901
Family Rhamphognathidae
Genus Rhamphognathus Agassiz, 1844
Rhamphognathus paralepoides Agassiz, 1844
Family Mesogasteridae
Genus Latellagnathus Bannikov, 2008
Latellagnathus teruzzii Bannikov, 2008
Genus Mesogaster Agassiz, 1844
Mesogaster sphyraenoides Agassiz, 1844
Order Beloniformes
Family Exocoetidae
Genus Rhamphexocoetus Bannikov, Parin et Pinna,
1985
Rhamphexocoetus volans Bannikov, Parin et Pinna, 1985
Family Hemiramphidae
Genus Hemiramphus Cuvier, 1817
Hemiramphus edwardsi Bassani, 1876
Beloniformes inc. sed.
“Engraulis” evolans (Blainville, 1818)
Series Percomorpha
Order Beryciformes
Family Holocentridae
Genus Berybolcensis Sorbini, 1984
Berybolcensis leptacanthus (Agassiz, 1838)
Genus Eoholocentrum Sorbini et Tirapelle, 1975
The systematic composition of the Eocene actinopterygian fish fauna from Monte Bolca, northern Italy
Eoholocentrum macrocephalum (Blainville, 1818)
(=Pygaeus concinnus Leriche, 1906)
Genus Tenuicentrum Sorbini, 1975
Tenuicentrum lanceolatum (Bassani, 1876)
Order Syngnathiformes
Suborder Syngnathoidei
Superfamily Pegasoidea
Family Rhamphosidae
Genus Rhamphosus Agassiz, 1839
Rhamphosus rastrum (Volta, 1796)
Rhamphosus biserratus Bassani, 1876
Superfamily Aulostomoidea
Family Urosphenidae
Genus Urosphen Agassiz, 1839
Urosphen dubius (Blainville, 1818)
Family Aulostomidae
Genus Eoaulostomus Blot, 1980
Eoaulostomus bolcensis (Blainville, 1818)
Eoaulostomus gracilis Blot, 1980
Genus Synhypuralis Blot, 1980
Synhypuralis jungerseni Blot, 1980
Synhypuralis banisteri Blot, 1980
Genus Jungensenichthys Blot, 1980
Jungensenichthys elongatus Blot, 1980
Genus Macraulostomus Blot, 1980
Macraulostomus veronensis Blot, 1980
Genus Tyleria N.N.Parin, 1993
Tyleria necopinata N.N.Parin, 1993
Family Parasynarcualidae
Genus Parasynarcualis Blot, 1980
Parasynarcualis longirostris (Blainville, 1818)
Family Fistularioididae
Genus Fistularioides Blot, 1980
Fistularioides veronensis Blot, 1980
Fistularioides phyllolepis Blot, 1980
Genus Pseudosyngnathus Kner et Steindachner, 1863
Pseudosyngnathus opisthopterus (Agassiz, 1833)
Aulostomoidea inc. sed.
Genus Aulostomoides Blot, 1980
Aulostomoides tyleri Blot, 1980
Superfamily Centriscoidea
Family Aulorhamphidae
Genus Aulorhamphus de Zigno, 1887
Aulorhamphus bolcensis (Steindachner, 1863)
Aulorhamphus capellinii Zigno, 1887
Aulorhamphus chiarasorbiniae Bannikov et Tyler, 2011
Genus Veronarhamphus Tyler, 2004
Veronarhamphus canossae (Heckel, 1856)
Genus Pesciarhamphus Bannikov et Tyler, 2011
Pesciarhamphus carnevalei Bannikov et Tyler, 2011
Family Paraeoliscidae
Genus Paraeoliscus Blot, 1980
Paraeoliscus robinetae Blot, 1980
Family Centriscidae
Genus Aeoliscoides Blot, 1980
Aeoliscoides longirostris (Blainville, 1818)
Genus Paramphisile Blot, 1980
Paramphisile weleri Blot, 1980
Superfamily Syngnathoidei
Family Syngnathidae
Genus “Syngnathus”
“Syngnathus” heckeli de Zigno, 1874
“Syngnathus” bolcensis de Zigno, 1887
Genus Prosolenostomus Blot, 1980
Prosolenostomus lessinii Blot, 1980
Family Solenostomidae (?)
Genus Solenorhynchus Heckel, 1854
Solenorhynchus elegans Heckel, 1854
Genus Calamostoma Agassiz, 1833
Calamostoma breviculum (Blainville, 1818)
Order Dactylopteriformes
Family Pterygocephalidae
Genus Pterygocephalus Agassiz, 1839
Pterygocephalus paradoxus Agassiz, 1839
Order Perciformes
Suborder Percoidei
Family Latidae
Genus Eolates Sorbini, 1970
Eolates gracilis (Agassiz, 1833)
Family Percichthyidae
Genus Cyclopoma Agassiz, 1833
Cyclopoma gigas Agassiz, 1833
Family Acropomatidae
Genus Acropoma Temminck et Schlegel, 1843
Acropoma lepidotum (Agassiz, 1836)
Family Priacanthidae
Genus Pristigenys Agassiz, 1839
Pristigenys substriatus (Blainville, 1818)
Family Apogonidae
Genus Eosphaeramia Sorbini, 1983
Eosphaeramia pygopterus (Agassiz, 1836)
Genus Eoapogon Bannikov, 2005
Eoapogon fraseri Bannikov, 2005
Genus Bolcapogon Bannikov, 2005
Bolcapogon johnsoni Bannikov, 2005
Genus Apogoniscus Bannikov, 2005
Apogoniscus pauciradiatus Bannikov, 2005
Apogonidae gen. et sp. indet.
Family Pomatomidae
Genus Carangopsis Agassiz, 1844
Carangopsis brevis (Blainville, 1818)
Carangopsis dorsalis Agassiz, 1844
Family Ductoridae
Genus Ductor Agassiz, 1844
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ALEXANDRE F. BANNIKOV
Ductor vestenae (Volta, 1796)
Family Carangidae
Genus Seriola Cuvier, 1817
Seriola prisca (Agassiz, 1834)
Genus Vomeropsis Heckel, 1854
Vomeropsis triurus (Volta, 1796)
Genus Ceratoichthys Blot, 1969
Ceratoichthys pinnatiformis (Blainville, 1818)
Genus Eastmanalepes Bannikov, 1984
Eastmanalepes primaevus (Eastman, 1904)
Genus Lichia Cuvier, 1817
Lichia veronensis Bannikov et Sorbini in Bannikov,
1990
Genus Paratrachinotus Blot, 1969
Paratrachinotus tenuiceps (Agassiz, 1834)
Family Apolectidae
Genus Trachicaranx Daniltshenko, 1968
Trachicaranx pleuronectiformis (Blot, 1969)
Family Menidae
Genus Mene Lacépède, 1803
Mene rhombea (Volta, 1796)
Mene oblonga (Agassiz, 1833)
Family Leiognathidae
Genus Eoleiognathus Bannikov, 2014
Eoleiognathus dorsalis (Agassiz, 1838)
Family Exelliidae
Genus Exellia White et Moy-Thomas, 1941
Exellia velifer (Volta, 1796)
Family Lutjanidae
Genus Ottaviania Sorbini, 1983
Ottaviania mariae Sorbini, 1983
Ottaviania leptacanthus (Agassiz, 1839)
Genus Veranichthys Bannikov, 2006
Veranichthys ventralis (Agassiz, 1839)
Genus Goujetia Bannikov, 2006
Goujetia crassispina (Agassiz, 1839)
Genus Lessinia Bannikov et Zorzin, 2014
Lessinia horrenda Bannikov et Zorzin, 2014
Lessinia sp.
Family Gerreidae (?)
Genus Aspesiperca Bannikov, 2008
Aspesiperca ruffoi Bannikov, 2008
Family Sparidae
Genus Sparnodus Agassiz, 1838
Sparnodus elongatus Agassiz, 1839
Sparnodus vulgaris (Blainville, 1818)
Genus Pseudosparnodus Day, 2003
Pseudosparnodus microstomus (Agassiz, 1839)
Genus Ellaserrata Day, 2003
Ellaserrata monksi Day, 2003
Genus “Dentex”
“Dentex” microdon Agassiz, 1839
Genus Abromasta Day, 2003
Abromasta microdon (Agassiz, 1839)
Nomen dubium:
“Dentex” ventralis Agassiz, 1839
Family Quasimullidae
Genus Quasimullus Bannikov, 1999
Quasimullus sorbinii Bannikov, 1999
Family Monodactylidae
Genus Psettopsis Blot, 1969
Psettopsis subarcuatus (Blainville, 1818)
Psettopsis latellai Bannikov, 2005
Family Ephippididae
Genus Archaephippus Blot, 1969
Archaephippus asper (Volta, 1796)
Family Platacidae
Genus Eoplatax Blot, 1969
Eoplatax papilio (Volta, 1796)
Family Scatophagidae
Genus Eoscatophagus Tyler et C.Sorbini, 1999
Eoscatophagus frontalis (Agassiz, 1839)
Family Pomacentridae
Genus Palaeopomacentrus Bellwood et Sorbini,
1996
Palaeopomacentrus orphae Bellwood et Sorbini, 1996
Genus Lorenzichthys Bellwood, 1999
Lorenzichthys olihan Bellwood, 1999
Genus Sorbinichromis Bannikov et Bellwood, 2014
Sorbinichromis francescoi Bannikov et Bellwood, 2014
Family Carangodidae
Genus Carangodes Heckel, 1856
Carangodes bicornis (Volta, 1796) (C. cephalus =Heckel,
1856; syn. nov.)
Family Eocottidae
Genus Eocottus Woodward, 1901
Eocottus veronensis (Volta, 1796)
Genus Bassanichthys Bannikov, 2006
Bassanichthys pesciaraensis (Bannikov, 2004)
Family Robertanniidae
Genus Robertannia Bannikov, 2011
Robertannia sorbiniorum Bannikov, 2011
Genus Hendrixella Bannikov et Carnevale, 2009
Hendrixella grandei Bannikov et Carnevale, 2009
Percoidei inc. sed.
Genus Veronabrax Bannikov, 2008
Veronabrax schizurus (Agassiz, 1836)
Genus Voltamulloides Bannikov, 2008
Voltamulloides ceratorum Bannikov, 2008
Genus Parapelates Bannikov, 2008
Parapelates quindecimalis (Agassiz, 1836)
Genus Jimtylerius Bannikov et Carnevale, 2007
Jimtylerius temnopterus (Agassiz, 1836)
Genus Pavarottia Bannikov et Zorzin, 2011
The systematic composition of the Eocene actinopterygian fish fauna from Monte Bolca, northern Italy
Pavarottia lonardonii Bannikov et Zorzin, 2011
Genus Montepostalia Bannikov et Zorzin, 2004
Montepostalia annamariae Bannikov et Zorzin, 2004
Genus Blotichthys Sorbini, 1984
Blotichthys coleanus (Agassiz, 1838)
Genus Pygaeus Agassiz, 1838
Pygaeus bolcanus (Volta, 1796)
Pygaeus nobilis Agassiz, 1838
Pygaeus nuchalis Agassiz, 1838
Genus Malacopygaeus Leriche, 1906
Malacopygaeus oblongus (Agassiz, 1838)
Genus Gillidia Eastman, 1914
Gillidia antiqua (Agassiz, 1835)
Genus Bradyurus Gill, 1904
Bradyurus szajnochae (de Zigno, 1887)
Genus Frigoichthys Bannikov, 2004
Frigoichthys margaritae Bannikov, 2004
Genus Squamibolcoides Bannikov et Zorzin, 2013
Squamibolcoides minciottii Bannikov et Zorzin, 2013
Genus Frippia Bannikov et Carnevale, 2012
Frippia labroiformis Bannikov et Carnevale, 2012
Suborder Sphyraenoidei
Family Sphyraenidae
Genus Sphyraena Bloch et Schneider, 1801
Sphyraena bolcensis Agassiz, 1844
Suborder Labroidei
Family Tortonesidae
Genus Tortonesia Sorbini, 1983
Tortonesia esilis Sorbini, 1983
Family Labridae
Genus Eocoris Bannikov et Sorbini, 1990
Eocoris bloti Bannikov et Sorbini, 1990
Genus Phyllopharyngodon Bellwood, 1990
Phyllopharyngodon longipinnis Bellwood, 1990
Genus Bellwoodilabrus Bannikov et Carnevale,
2010
Bellwoodilabrus landinii Bannikov et Carnevale, 2010
Labroidei inc. sed.
Genus “Labrus”
“Labrus” valenciennesi Agassiz, 1839
Genus Sorbinia Bellwood, 1995
Sorbinia caudopunctata Bellwood, 1995
Suborder Trachinoidei
Family Callipterygidae
Genus Callipteryx Agassiz, 1838
Callipteryx recticaudus Agassiz, 1838
Callipteryx speciosus Agassiz, 1838
Suborder Gobioidei
Gobioidei (?) inc. sed.
Genus “Gobius”
“Gobius” microcephalus Agassiz, 1839
Suborder Acanthuroidei
29
Family Caproidae
Genus Eoantigonia Baciu, Bannikov et Tyler, 2005
Eoantigonia veronensis (Sorbini, 1983)
Family Sorbinipercidae
Genus Sorbiniperca Tyler, 1999
Sorbiniperca scheuchzeri Tyler, 1999
Genus Sorbinicapros Bannikov et Tyler, 1999
Sorbinicapros sorbiniorum Bannikov et Tyler, 1999
Family Zorzinichthyidae
Genus Zorzinichthys Tyler et Bannikov, 2002
Zorzinichthys annae Tyler et Bannikov, 2002
Family Acanthonemidae
Genus Acanthonemus Agassiz, 1834
Acanthonemus subaureus (Blainville, 1818)
Family Siganidae
Genus Ruffoichthys Sorbini, 1983
Ruffoichthys spinosus Sorbini, 1983
Ruffoichthys bannikovi Tyler et Sorbini, 1990
Genus Aspesiganus Bannikov et Tyler, 2002
Aspesiganus margaritae Bannikov et Tyler, 2002
Genus Acanthopygaeus Leriche, 1906
Acanthopygaeus agassizi (Eastman, 1904)
Family Acanthuridae
Genus Proacanthurus Blot et Tyler, 1990
Proacanthurus tenuis (Agassiz, 1838)
Proacanthurus bonatoi Blot et Tyler, 1990
Proacanthurus ovalis (Agassiz, 1838)
Proacanthurus elongatus Blot et Tyler, 1990
Genus Metacanthurus Blot et Tyler, 1990
Metacanthurus veronensis Blot et Tyler, 1990
Genus Eorandallius Blot et Tyler, 1990
Eorandallius rectifrons (Agassiz, 1838)
Eorandallius elegans Blot et Tyler, 1990
Genus Acanthuroides Blot et Tyler, 1990
Acanthuroides massalongoi Blot et Tyler, 1990
Genus Lehmanichthys Blot et Tyler, 1990
Lehmanichthys lessiniensis Blot et Tyler, 1990
Genus Metaspisurus Blot et Tyler, 1990
Metaspisurus emmanueli Blot et Tyler, 1990
Genus Pesciarichthys Blot et Tyler, 1990
Pesciarichthys punctatus Blot et Tyler, 1990
Genus Frigosorbinia Bannikov et Tyler, 2013
Frigosorbinia baldwinae (Sorbini et Tyler, 1998)
Genus Tylerichthys Blot, 1980
Tylerichthys nuchalis (Agassiz, 1838)
Tylerichthys milani Blot et Tyler, 1990
Genus Protozebrasoma Sorbini et Tyler, 1998
Protozebrasoma bloti Sorbini et Tyler, 1998
Genus Sorbinithurus Tyler, 1999
Sorbinithurus sorbinii Tyler, 1999
Genus Tauichthys Tyler, 1999
Tauichthys padremenini Tyler, 1999
30
ALEXANDRE F. BANNIKOV
Tauichthys aspesae Tyler et Bannikov, 2000
Genus Gazolaichthys Blot et Tyler, 1990
Gazolaichthys vestenanovae Blot et Tyler, 1990
Genus Padovathurus Tyler, 2005
Padovathurus gaudryi (de Zigno, 1887)
Family Zanclidae
Genus Eozanclus Blot et Voruz, 1975
Eozanclus brevirostris (Agassiz, 1835)
Family Massalongiidae
Genus Massalongius Tyler et Bannikov, 2005
Massalongius gazolai (Massalongo, 1859)
Suborder Scombroidei
Family Euzaphlegidae
Genus Veronaphleges Bannikov, 2008
Veronaphleges brunae Bannikov, 2008
Family Scombridae
Genus Auxides Jordan, 1919
Auxides propterygius (Agassiz, 1835)
Genus Pseudauxides Monsch, 2006
Pseudauxides speciosus (Agassiz, 1835)
Genus Thunnoscomberoides Monsch, 2006
Thunnoscomberoides bolcensis (Agassiz, 1835)
Genus Godsilia Monsch, 2006
Godsilia lanceolata (Agassiz, 1835)
Suborder Xiphioidei
Family Blochiidae
Genus Blochius Volta, 1796
Blochius longirostris Volta, 1796
Blochius macropterus de Zigno, 1887
Family Palaeorhynchidae
Genus Palaeorhynchus Blainville, 1818
Palaeorhynchus zorzini Fierstine, Bannikov et Monsch,
2008
Suborder Stromateoidei
Family Centrolophidae
Genus Zorzinia Bannikov, 2000
Zorzinia postalensis Bannikov, 2000
Perciformes inc. sed.
Genus Quasicichla Bannikov, 2004
Quasicichla mucistonaver Bannikov, 2004
Genus Parapygaeus Pellégrin, 1907
Parapygaeus polyacanthus Pellégrin, 1907
Order Pleuronectiformes
Family Amphistiidae
Genus Amphistium Agassiz, 1844
Amphistium paradoxum Agassiz, 1844
Genus Heteronectes Friedman, 2008
Heteronectes chaneti Friedman, 2008
Pleuronectiformes inc. sed.
Genus Eobothus Eastman, 1914
Eobothus minimus (Agassiz, 1839)
Order Tetraodontiformes
Suborder Triacanthoidei
Family Triacanthidae
Genus Protacanthodes Gill, 1888
Protacanthodes ombonii (de Zigno, 1887)
Protacanthodes nimesensis Tyler et Santini, 2001
Suborder Balistoidei
Family Protobalistidae
Genus Spinacanthus Agassiz, 1844
Spinacanthus cuneiformis (Blainville, 1818)
Genus Protobalistum de Zigno, 1887
Protobalistum imperiale (Massalongo, 1857)
Family Bolcabalistidae
Genus Bolcabalistes Tyler et Sorbini, 1998
Bolcabalistes varii Tyler et Sorbini, 1998
Family Aracanidae
Genus Proaracana Le Danois, 1961
Proaracana dubia (Blainville, 1818)
Family Ostraciidae
Genus Eolactoria Tyler, 1975
Eolactoria sorbinii Tyler, 1975
Suborder Tetraodontoidei
Family Eoplectidae
Genus Eoplectus Tyler, 1975
Eoplectus bloti Tyler, 1975
Family Zignoichthyidae
Genus Zignoichthys Tyler, 1975
Zignoichthys oblongus (de Zigno, 1874)
Family Tetraodontidae
Genus Eotetraodon Tyler, 1980
Eotetraodon pygmaeus (de Zigno, 1887)
Eotetraodon tavernei Tyler et Bannikov, 2013
Family Diodontidae
Genus Prodiodon Le Danois, 1955
Prodiodon tenuispinus (Agassiz, 1844)
Prodiodon erinaceus (Agassiz, 1844)
Genus Heptadiodon Bronn, 1855
Heptadiodon echinus (Heckel, 1854)
Genus Zignodon Tyler et Santini, 2002
Zignodon fornasieroae Tyler et Santini, 2002
Acanthopterygii inc. sed.
Genus Pietschellus Bannikov et Carnevale, 2011
Pietschellus aenigmaticus Bannikov et Carnevale, 2011
Genus Xiphopterus Agassiz, 1844
Xiphopterus falcatus (Volta, 1796)
Genus Oncolepis Bassani, 1897
Oncolepis isseli Bassani, 1897
Genus Protaulopsis Woodward, 1901
Protaulopsis bolcensis Woodward, 1901
The systematic composition of the Eocene actinopterygian fish fauna from Monte Bolca, northern Italy
Thus, according to the latest data, the known Actinopterygii in the Bolca fauna belong to 19 orders, 91
families, 192 genera (32 of which are of uncertain family) and at least 220 species; among these, the Acanthomorpha represent 11 orders, 78 families, 163 genera and
at least 190 species. Undoubtedly this picture of the already amazing diversity of the Monte Bolca fish assemblage will be enhanced by future investigations. There
are still new taxa to be described, and some known taxa
are in need of extensive revision.
While in the list of Bolca fishes published by Blot
(1980) most of the teleostean families are extant (56, including three doubtful), and only 15 families are extinct
(systematic position of 10 genera, including four undescribed, uncertain), subsequent investigations have reduced the number of the extant families to 52 (including questionable Solenostomidae and Gerreidae). At
the same time, the number of the extinct families in
the Monte Bolca fish fauna has increased to 39, whereas
the systematic positions of 32 genera remain uncertain;
most of the latter belong to fossil families.
Acknowledgements
I am grateful to Dr. Giuseppe Minciotti, Director of
the Museo Civico di Storia Naturale di Verona, for providing funding for my travel in 2014 to Verona to engage in this and other studies. I also thank: Dr. Roberto Zorzin and Dr. Anna Vaccari at the MCSNV for
their continuing help in facilitating our research on the
fishes of Monte Bolca; Dr. Leonardo Latella for editorial processing of the manuscript; and Mrs. Bruna Burato for her aid with the literature on Bolca fishes. Dr.
James C. Tyler of the Smithsonian Institution kindly
reviewed the manuscript and improved the English,
and Dr. Chiara Sorbini of the University of Pisa translated the Abstract into Italian. My research was supported by the Russian Foundation for Basic Research,
grant no. 14-04-00005.
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Address of the author
ALEXANDRE F. BANNIKOV
Borisyak Paleontological Institute of the Russian Academy of Sciences,
Profsoyuznaya 123, Moscow 117997, Russia
e-mail: aban@paleo.ru

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