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 24 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 25 26 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 27 28 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. Literature cited AGASSIZ L., 1833-1844. Recherches sur les poissons fossiles. Petitpierre, Neuchâtel: Tome IV: 296 pp.; Tome V, pt 1: 122 pp. BANNIKOV A.F., 1991 (1990). An Eocene veliferoid (Teleostei, Lampridiformes) from Bolca. Miscellanea paleontologica. Studi e Ricerche sui Giacimenti Terziari di Bolca, 6: 161-174. BANNIKOV A.F., 1999. Unusual new percoid fish 31 from the Eocene of Bolca. 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British Museum (Natural History), London: 638 pp. ZORZIN R., BANNIKOV A.F., FORNACIARI E., GIUSBERTI L., PAPAZZONI C.A., ROGHI G., 2011. Il giacimento a pesci e piante fossili dell’Eocene Inferiore di Monte Solane (Verona). Bollettino del Museo Civico di Storia Naturale di Verona. Geologia Paleontologia Preistoria, 35: 57-64. Address of the author ALEXANDRE F. BANNIKOV Borisyak Paleontological Institute of the Russian Academy of Sciences, Profsoyuznaya 123, Moscow 117997, Russia e-mail: [email protected]
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