Fig 1.
Sketch map of northeastern Italy showing the location of the Bolca, Monte Solane, and Solteri Lagerstätten.
Scale bar 50 km. Map modified and reprinted from [14] under a CC BY license, with permission from Rivista Italiana di Paleontologia e Stratigrafia, original copyright year 2025.
Fig 2.
Correlation between the Ypresian Monte Solane (Verona) and Solteri (Trento) fish-bearing Lagerstätten.
A: chronostratigraphy; B: larger foraminiferal zonation SB after Serra-Kiel et al. [42]; C: calcareous nannofossil zonation CNE after Agnini et al. [41] and equivalent zonation NP of Martini [43]; D: planktic foraminiferal zonation E after Wade et al. [28], modified by [29]. Lithologic legend: 1) laminated, organic-rich marls and calcareous marls; 2) limestones, marly limestones and marls; 3) calcarenitic limestones; 4) calciruditic larger foraminiferal limestones; 5) chert; 6) dark grey to brown lithologies; 7) fishes.
Fig 3.
Single isolated tooth exposed on the labial side. Scale bar 10 mm.
Table 1.
Measurements of Bolcaichthys solanensis n. sp. compared to the type species of the genus B. catopygopterus.
Fig 4.
Bolcaichthys solanensis n. sp.
Lateral view of the holotype, MGP-PD 33556, in natural light (A) and coated in alcohol (B-D). Details of the cranium (C) and of the caudal fin (D). Scale bars 10 mm (A-B); 5 mm (C-D).
Fig 5.
Bolcaichthys solanensis n. sp.
Holotype, MGP-PD 33556, interpretive reconstruction of the cranium. Scale bar 5 mm. Abbreviations: aa: anguloarticular; ao: antorbital; achy: anterior ceratohyal; br: branchiostegal rays; bsph: basisphenoid; cor: coracoid; cl: cleithrum; den: dentary; ect: ectopterygoid; end: endopterygoid; fr: frontal; hyo: hyomandibula; IO: infraorbital bones; let: lateral ethmoid; mes: mesethmoid; mtp: metapterygoid; mx: maxilla; op: opercle; osph: orbitosphenoid; pa: parietal; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pts: pterosphenoid; ptt: posttemporal; q: quadrate; sca: scapula; scl: supracleithrum; so: supraorbital; soc: supraoccipital; sop: subopercle; sph: sphenotic; sr: sclerotic ring; sym: symplectic; tf: temporal fossa; vom: vomer.
Fig 6.
Bolcaichthys solanensis n. sp.
Holotype, MGP-PD 33556, interpretive reconstruction of the caudal skeleton. Scale bar 5 mm. Abbreviations: ep: epural; h: hypural; hspu2: haemal spine of the second preural vertebra; hspu3: haemal spine of the third preural vertebra; nspu2: neural spine of the second preural vertebra; nspu3: neural spine of the third preural vertebra; ph: parhypural; pu1: first preural centrum; u1: first ural centrum; un: uroneural.
Table 2.
Summary of the meristic traits of the different species of the genus Bolcaichthys.
Fig 7.
Lepidoclupea renga n. gen. et n. sp.
Lateral view of the holotype, IGVR 64083-67873 (A-B). Lateral view of the paratypes, coated in alcohol, MGP-PD 33506 (C), MGP-PD 33517 (D), MGP-PD 33535 (E), MGP-PD 33528 (F). Scale bars 20 mm.
Fig 8.
Lepidoclupea renga n. gen. et n. sp. Holotype.
Interpretive reconstruction of the cranium. IGVR 64083 (A), IGVR 67873 (B). Scale bars 5 mm. Abbreviations: aa: anguloarticular; achy: anterior ceratohyal; br: branchiostegal rays; cb: ceratobranchial; cl: cleithrum; den: dentary; ect: ectopterygoid; end: endopterygoid; fr: frontal; hyo: hyomandibula; IO: infraorbital; let: lateral ethmoid; mtp: metapterygoid; mx: maxilla; op: opercle; pa: parietal; pal: palatine; pchy: posterior ceratohyal; pcl: postcleithrum; pfr: pectoral-fin ray; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pts: pterosphenoid; q: quadrate; rad: pectoral-fin radial; sca: scapula; scl: supracleithrum; sop: subopercle; sph: sphenotic; sym: symplectic; vom: vomer.
Table 3.
Measurements of Lepidoclupea renga n. gen. et n. sp. compared with the Eocene round herring Trollichthys bolcensis.
Table 4.
Summary of the meristic traits of Lepidoclupea renga n. gen. et n. sp. compared to those of the round herring Trollichthys bolcensis.
Table 5.
Measurements of Sabbathichthys osbournei n. gen. et n. sp.
Table 6.
Summary of the meristic traits of Sabbathichthys osbournei n. gen. et. n. sp. and other extinct photichthyid taxa.
Table 7.
Measurements of the different species of the genus Scopeloides.
Table 8.
Summary of the meristic traits of the different species of the genus Scopeloides.
Table 10.
Summary of the meristic traits of the different species of the genus Eomyctophum and Oligophus moravicus.
Table 9.
Measurements of Eomyctophum mainardii from Monte Solane.
Fig 9.
Lepidoclupea renga n. gen. et n. sp.
Holotype, IGVR 64083. Interpretive reconstruction of the caudal skeleton (A) and pelvic girdle (B). Scale bars 5 mm (A); 1 mm (B). Abbreviations: bas: basipterygium; ep: epural; h: hypural; hspu2: haemal spine of the second preural vertebra; hspu3: haemal spine of the third preural vertebra; nspu2: neural spine of the second preural vertebra; nspu3: neural spine of the third preural vertebra; ph: parhypural; pu1: first preural centrum; scu: abdominal scute; u1: first ural centrum; u2: second ural centrum; vfr: pelvic fin rays.
Fig 10.
Sabbathichthys osbournei n. gen. et n. sp.
Lateral view of the holotype, MGP-PD 33551, in natural light (A) and coated in alcohol (B). Lateral view of the paratype, IGVR 64077, in natural light (C) and coated in alcohol (D). Interpretive reconstruction of the skeleton of the holotype (E). Scale bars 10 mm.
Fig 11.
Sabbathichthys osbournei n. gen. et n. sp.
Holotype, MGP-PD 33551. Interpretive reconstruction of the cranium. Scale bar 10 mm. Abbreviations: aa: anguloarticular; br: branchiostegal rays; cb: ceratobranchial; cor: coracoid; cl: cleithrum; den: dentary; ect: ectopterygoid; end: endopterygoid; fr: frontal; hyo: hyomandibula; mtp: metapterygoid; mx: maxilla; op: opercle; pa: parietal; pal: palatine; pfr: pectoral-fin ray; pmx: premaxilla; pop: preopercle; ptt: posttemporal; q: quadrate; rad: pectoral-fin radial; scl: supracleithrum; uhy: urohyal.
Fig 12.
Sabbathichthys osbournei n. gen. et n. sp.
Holotype, MGP-PD 33551. Interpretive reconstruction of the caudal skeleton. Scale bar 5 mm. Abbreviations: ep: epural; h: hypural; hspu2: haemal spine of the second preural vertebra; hspu3: haemal spine of the third preural vertebra; nspu2: neural spine of the second preural vertebra; nspu3: neural spine of the third preural vertebra; ph: parhypural; un: uroneural; ur: urostyle.
Fig 13.
Scopeloides bellator Calzoni, Giusberti & Carnevale, 2025 [14]. Lateral view of IGVR 64073-67862 (A-B), IGVR 64074 (C), IGVR 82447 (D), IGVR 64075-82438 (E-F), MGP-PD 33457 (G), MGP-PD 33500 (H). Scale bars 10 mm.
Fig 14.
Scopeloides violator Calzoni, Giusberti & Carnevale, 2025 [14]. Lateral view of MGP-PD 33470 (A), MGP-PD 33479 (B); MGP-PD 33504 (C), MGP-PD 33510 (D). Scale bars 10 mm.
Fig 15.
Lateral view of IGVR 82433-82434 (A-B), MGP-PD 33451A-B (C-D), MGP-PD 33463A-B (E-F), MGP-PD 33550 (H), and dorsal view of MGP-PD 33503 (G), all coated in alcohol. Scale bars 10 mm.
Fig 16.
Eomyctophum mainardii Calzoni, Giusberti & Carnevale, 2025 [14]. Lateral view of IGVR 64078-67861 (A-B), IGVR 67863 (C), IGVR 67856b (D). Scale bars 10 mm.
Fig 17.
Acanthophleges lessiniae n. gen. et n. sp.
Lateral view of the holotype, IGVR 64085-67877, in normal light (A-B) and detail of the cranium coated in alcohol (C-D). Interpretive reconstruction of the cranium (E). Scale bars 10 mm (A-D); 5 mm (E). Abbreviations: aa: anguloarticular; bas: basipterygium; br: branchiostegal rays; bsph: basisphenoid; cl: cleithrum; cor: coracoid; den: dentary; end: endopterygoid; epi: epioccipital; exsc: extrascapular; fr: frontal; hyo: hyomandibula; let: lateral ethmoid; mes: mesethmoid; mx: maxilla; op: opercle; pa: parietal; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pto: pterotic; pts: pterosphenoid; ptt: posttemporal; q: quadrate; sca: scapula; scl: supracleithrum; soc: supraoccipital; sop: subopercle; sph: sphenotic; vom: vomer; vpcl: ventral postcleithrum.
Table 11.
Summary of the meristic traits of different euzaphlegid taxa.
Fig 18.
Lateral view of the holotype, IGVR 64084 (A). Detail of the cranium, coated in alcohol (B). Interpretive reconstruction of the cranium (C). Scale bars 20 mm (A); 10 mm (B-C). Abbreviations: aa: anguloarticular; achy: anterior ceratohyal; br: branchiostegal rays; den: dentary; ect: ectopterygoid; end: endopterygoid; fr: frontal; hh: ventral hypohyal; hyo: hyomandibula; iop: interopercle; la: lachrymal; let: lateral ethmoid; mes: mesethmoid; mtp: metapterygoid; mx: maxilla; op: opercle; pa: parietal; pal: palatine; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pto: pterotic; pts: pterosphenoid; ptt: posttemporal; q: quadrate; soc: supraoccipital; sop: subopercle; sph: sphenotic; sym: symplectic; vom: vomer.
Fig 19.
Holotype, IGVR 64084. Interpretive reconstruction of the skeleton. Scale bar 20 mm.
Table 12.
Measurements of Veronaphleges ambrosii n. sp. compared to the type species of the genus, Veronaphleges brunae.
Table 13.
Summary of the meristic traits of different gempylid taxa from Monte Solane.
Table 14.
Measurements of the various gempylid taxa from Monte Solane.
Fig 20.
Contemptor mastinoi n. gen. et n. sp.
Lateral view in natural light of the holotype, IGVR 82435-82436 (A-B), detail of the cranium of IGVR 82345 coated in alcohol (C). Lateral view of the paratype, IGVR 67858, coated in alcohol (D) and of the specimen IGVR 64080a-b in natural light (E-F). Scale bars 20 mm (A-B, D-F), 10 mm (C).
Fig 21.
Contemptor mastinoi n. gen. et n. sp.
Holotype, IGVR 82435-82436. Interpretive reconstruction of the cranium. Scale bar 10 mm. Abbreviations: aa: anguloarticular; achy: anterior ceratohyal; bas: basipterygium; br: branchiostegal rays; bsph: basisphenoid; cb: ceratobranchial; cl: cleithrum; cor: coracoid; den: dentary; ect: ectopterygoid; end: endopterygoid; fr: frontal; hyo: hyomandibula; IO2: second infraorbital bone; iop: interopercle; la; lachrymal; let: lateral ethmoid; mes: mesethmoid; mtp: metapterygoid; mx: maxilla; op: opercle; pa: parietal; pal: palatine; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pto: pterotic; pts: pterosphenoid; ptt: posttemporal; q: quadrate; sca: scapula; scl: supracleithrum; smx: supramaxilla; soc: supraoccipital; sop: subopercle; sph: sphenotic; spn: supraneural; sym: symplectic; vom: vomer; vpcl: ventral postcleithrum.
Fig 22.
Contemptor mastinoi n. gen. et n. sp.
Interpretive reconstruction of the caudal skeleton. Scale bar 5 mm. Abbreviations: ep: epural; h: hypural; hspu2: haemal spine of the second preural vertebra; hspu3: haemal spine of the third preural vertebra; nspu2: neural spine of the second preural vertebra; nspu3: neural spine of the third preural vertebra; ph: parhypural; u: ural centrum.
Fig 23.
Krampusichthys tridentinus Calzoni, Giusberti & Carnevale, 2025 [14].
Lateral view of IGVR 64109 (A), MGP-PD 33459 (B), and MGP-PD 33511 (C). Scale bars 10 mm.
Fig 24.
Thyrsitoides cangrandei n. sp.
Lateral view in natural light and coated in alcohol of the holotype, IGVR 82445 (A). Lateral view of the paratype IGVR 67880, coated in alcohol and detail of the cranium (B-C). Interpretive reconstruction of the skeleton (infraorbital series is omitted; D). Scale bars 10 mm.
Fig 25.
Thyrsitoides cangrandei n. sp.
Paratype, IGVR 67780. Interpretive reconstruction of the cranium. Scale bar 10 mm. Abbreviations: aa: anguloarticular; achy: anterior ceratohyal; bas: basipterygium; br: branchiostegal rays; bsph: basisphenoid; cl: cleithrum; cor: coracoid; den: dentary; ect: ectopterygoid; end: endopterygoid; epi:epioccipital; fr: frontal; hyo: hyomandibula; IO2-3: infraorbital bones; iop: interopercle; la; lachrymal; let: lateral ethmoid; mes: mesethmoid; mtp: metapterygoid; mx: maxilla; na: nasal; op: opercle; pa: parietal; pchy: posterior ceratohyal; pfr: pectoral-fin rays; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pto: pterotic; pts: pterosphenoid; ptt: posttemporal; q: quadrate; scl: supracleithrum; soc: supraoccipital; sop: subopercle; sph: sphenotic; spn: supraneural; sym: symplectic; vom: vomer; vfr: pelvic-fin rays; vpcl: ventral postcleithrum.
Table 15.
Measurements of the various species of the genus Thyrsitoides.
Table 16.
Summary of the meristic traits of different Thyrsitoides species.
Fig 26.
Thyrsitoides cangrandei n. sp.
Holotype, IGVR 82445. Interpretive reconstruction of the caudal skeleton. Scale bar 10 mm. Abbreviations: h: hypural; hspu2: haemal spine of the second preural vertebra; hspu3: haemal spine of the third preural vertebra; nspu2: neural spine of the second preural vertebra; nspu3: neural spine of the third preural vertebra; ph: parhypural; un: uroneural.
Fig 27.
Lateral view of MGP-PD 33557 in natural light (A) and coated in alcohol (B). Scale bars 20 mm.
Fig 28.
Eomastix zabimaru n. gen. et n. sp.
Lateral view of the holotype, IGVR 64081-64082, in natural light (A-B), IGVR 64082 coated in alcohol (C) and detail of the cranium of IGVR 64082 coated in alcohol (D). Interpretive reconstruction of the cranium (E). Scale bars 10 mm (A-D); 5 mm (E). Abbreviations: aa: anguloarticular; achy: anterior ceratohyal; bas: basipterygium; br: branchiostegal rays; cl: cleithrum; cor: coracoid; den: dentary; dpcl: dorsal postcleithrum; ect: ectopterygoid; end: endopterygoid; epi: epioccipital; fr: frontal; hh: ventral hypohyal; hyo: hyomandibula; iop: interopercle; let: lateral ethmoid; mes: mesethmoid; mtp: metapterygoid; mx: maxilla; op: opercle; pa: parietal; pchy: posterior ceratohyal; pmx: premaxilla; pop: preopercle; psp: parasphenoid; pto: pterotic; pts: pterosphenoid; ptt: posttemporal; q: quadrate; sca: scapula; scl: supracleithrum; smx: supramaxilla; soc: supraoccipital; sop: subopercle; sph: sphenotic; vfr: pelvic-fin rays; vom: vomer; vpcl: ventral postcleithrum.
Table 17.
Measurements of Eomastix zabimaru n. gen. et n. sp.
Table 18.
Summary of the meristic traits of Eomastix zabimaru n. gen. et n. sp. compared with different trichiurid taxa.
Fig 29.
Lateral view of IGVR 82413 in natural light (A) and coated in alcohol (B). Interpretive reconstruction of the skeleton of IGVR 82413. Scales are omitted (C). Scale bars 5 mm.
Table 19.
Meristic and characteristic traits of the Subfamily Apogoninae compared to IGVR 82413.
Table 20.
Summary of the morphological and meristic traits of IGVR 82413 compared with different genera of fossil apogonids.
Fig 30.
Comparison between the taxonomic composition of the ichthyofauna of the Monte Solane (A) and Solteri (B) sites.
Graph modified and reprinted from [14] under a CC BY license, with permission from Rivista Italiana di Paleontologia e Stratigrafia, original copyright year 2025.
Fig 31.
Biomagnetostratigraphy of the Ypresian p.p.-Lutetian p.p. interval.
Time scale, geomagnetic polarity scale, planktic foraminiferal zonation (E), larger benthic foraminiferal zonation (SB) and calcareous nannofossil zonations (NP and CE) after Speijer et al. [45] with modifications. Planktic foraminiferal zonation E of Wade et al. [28] with modifications of Luciani & Giusberti [29], Tethyan zonation of larger benthic foraminifera SB from Serra-Kiel et al. [42] with modifications of Benedetti et al. [142], and calcareous nannofossil zonations NP of Martini [43] and CNE of Agnini et al. [41]. The yellow bar corresponds to the EECO stratigraphical position and concept according to Westerhold et al. [141]. The blue arrow indicates the stratigraphic position of the Monte Solane and Solteri fish beds.