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Genus Libanopycnodus gen. nov. urn:lsid:zoobank.org:act:30B10E58-4147-4982-ADE0-B5956A502764 Type species Libanopycnodus wenzi gen. and sp. nov. (by monotypy). Diagnosis As for the species (monospecific genus). Etymology The generic eptithet refers to Lebanon and the taxon name Pycnodus is added., Published as part of Taverne, Louis & Capasso, Luigi, 2018, Osteology and relationships of Libanopycnodus wenzi gen. et sp. nov. and Sigmapycnodus giganteus gen. et sp. nov. (Pycnodontiformes) from the Late Cretaceous of Lebanon, pp. 1-29 in European Journal of Taxonomy 420 on page 4, DOI: 10.5852/ejt.2018.420, http://zenodo.org/record/1210447

Libanopycnodus wenzi gen. et sp. nov. urn:lsid:zoobank.org:act:2AD09A18-0416-4A26-AD08-E5307B901759 Figs 1���11 Diagnosis Small, deep-bodied pycnodontid fish, with rounded dorsal and ventral profiles. Head triangular in shape, with a rectilinear frontal profile and a short postorbital region. Pointed snout, with a mouth gape ventrally inclined. Frontal short. Prefrontal very narrow. Dorsal region of parietal forming a marked angle with the ventral region. Parietal brush-like process present. No temporal fenestra. Dermosphenotic sutured to the skull roof. Large, deep, well-visible dilatator fossa surrounded by dermosphenotic and dermopterotic. Posterior region of the endocranium exposed behind the dermopterotic. Vomer with 10 rounded molariform teeth on the external dental row. Premaxilla bearing two incisiform teeth. Prearticular with three rows of molariform teeth. Ectopterygoid present. Large triangular first infraorbital. Preopercle much deeper than the exposed region of the hyomandibula-dermohyomandibula. Opercle small and coma-shaped. Notochord not completely surrounded by vertebral arches. Neural and haemal spines with an anterior wing-like component. Twenty-five neural spines before epichordal series. First eight neural spines autogenous. Twelve haemal spines before hypochordal series. Eleven pairs of ribs. Postcoelomic bone long, narrow, with an enlarged ventral extremity. Dorsal and anal fins strip-like. Dorsal fin supported by 42 pterygiophores. Anal fin supported by 34 pterygiophores. Caudal peduncle short. Five epichordals. Eleven hypochordals, with two moderately broadened. One urodermal. Caudal fin double emarginated, with 20 principal rays. A few scale bars on the abdominal region of the body. Small complete scales associated with the prepelvic ventral keel scutes. Bifid cloacal scale present. One postcloacal scale. Dorsal ridge with 11 scutes, of which the first two with a small spine. Ventral keel with 13 scutes, 11 prepelvic and two postcloacal with small spines. Etymology The specific epithet is in homage to Dr. Sylvie Wenz (Paris), who worked abundantly on the pycnodontiform fishes. Material examined Holotype LEBANON: a complete specimen (Figs 1���2), total length: 90 mm, standard length: 78 mm, marine Upper Cenomanian deposits of Ein Namoura (CLC S-574). General morphology and morphometric data The fish is deep-bodied, the maximum body height being equal to a little more than the half of the standard length. The dorsal and the ventral profiles are rounded and devoid of marked apex. The morphometric data are given in % of the standard length (78 mm): Length of the head (opercle included) ���������������������������������������������������������������������34.0 % Depth of the head (in the occipital region) ���������������������������������������������������������������44.7 % Maximum depth of the body (just behind the head) ���������������������������������������������������56.2 % Prepelvic length ���������������������������������������������������������������������������������������������57.0 % Predorsal length ���������������������������������������������������������������������������������������������68.1 % Basal length of the dorsal fin ������������������������������������������������������������������������������35.7 % Preanal length ������������������������������������������������������������������������������������������������71.5 % Basal length of the anal fin ���������������������������������������������������������������������������������28.9 % Depth of the caudal peduncle ���������������������������������������������������������������������������������7.2 % Osteology Skull (Figs 3���6) The head is large, triangular in shape, altogether long and deep, with an almost rectilinear frontal profile. The preorbital region is much longer than the postorbital one. The orbit is wide. The dermal bones of the skull are ornamented with small tubercles. The snout is pointed, with a mouth gape ventrally inclined. The mesethmoid is elongate and broad. Its upper margin is covered posteriorly by a pair of very narrow prefrontals and anteriorly by the premaxillae. The orbitosphenoid is pressed against the posterior margin of the mesethmoid. The vomer, seen in profile, is long and deeper posteriorly than anteriorly. Only the lateral dental row, composed of 7 well-developed, rounded molariform teeth, is visible. There are traces of 3 other very small teeth located at the anterior extremity of the lower margin of the vomer. The frontal is short and limited to the orbital region. The dermosupraoccipital is small, with a pointed dorso-posterior extremity. The parietal exhibits an unusual and strange shape. Its dorsal half forms a marked angle with its ventral part. This peculiar shape of the parietal is not due to an artefact of fossilisation. A short brush-like process (= branched peniculus) is attached to the parietal. The supratemporal is a rather broad scale that lies against the upper part of the parietal, just below the first dorsal ridge scute. The dermopterotic is deeper than long. The dermosphenotic is not a free bone. It is included in the lateral wall of the skull roof and is sutured with the frontal and the dermopterotic. The ventral margin of the dermosphenotic and the dermopterotic are located at the same level as the lower border of the orbit. The dermopterotic and the dermosphenotic surround a deep and wide dilatator fossa. The parasphenoid is long, broad, toothless and its trabercular region is obliquely inclined. The posterior region of the endocranium is well visible. A small part of the intercalar and the exoccipital are preserved behind the lower half of the parietal. The ventral margin of the exoccipital is pierced by a large foramen for the branches of the vagus nerve (X). The basioccipital, articulated with the first basiventral, is also present. The exoccipital and basioccipital are separated from the braincase main part as in Pseudopycnodus nardoensis (Taverne 1997: fig. 2). A cartilaginous region probably linked the two cranial regions on the living fish. The autosphenotic and the sensory canals of the skull roof are not visible. A temporal fenestra is not present. The quadratic arch includes wide metapterygoid and entopterygoid plus a small ectopterygoid. Only the anterior parts of the quadrate and the symplectic are preserved. Both bones articulate with the lower jaw. The articular head of the symplectic is very broad. The premaxilla is long and narrow. It bears two incisiform teeth. The maxilla is lost. The posterior extremity of the dentary is preserved but not its anterior part with the teeth. The articular is small. The angular is not preserved. The prearticular is a large bone with a marked hook-like coronoid process, bearing three rows of molariform teeth. The dorsal row contains 6 teeth, the middle row 7 teeth and the ventral row at least 5 teeth. The teeth of the dorsal row are the smallest, with a slightly concave surface and their dorsal margin bears some little tubercles. The teeth of the lower row are the largest. They are deeper than long, with a slightly convex surface. The orbital series contains four bones. The first infraorbital is a large triangular bone. The second and third infraorbitals are tubular bones. As already written, the dermosphenotic is sutured to the skull roof. A few fragments of the sclerotic bony ring are visible just under the frontal. The exposed part of the hyomandibula-dermohyomandibula is much smaller than the preopercle to which it is sutured. The long ventral branch of the hyomandibula is covered by the preopercle but remains clearly visible. The upper margin of the hyomandibula bears a broad dorsal process that articulates in the dilatator fossa. The preopercle is deep and broad, with a rather narrow dorsal margin. The opercle is well developed, with an acuminate ventral tip and a broader upper part. The anterior ceratohyal and two branchiostegal rays are visible below the preopercle. Girdles (Figs 3���4) A small posttemporal is preserved behind the skull, at the level of the suture between the parietal and the dermopterotic. The hypercleithrum (= supracleithrum) is a long and thin bone. The cleithrum is divided in a short but very broad ventral branch and an extremely long but narrow dorsal branch. No pterygiophore is preserved but fragments of short pectoral rays are visible. The pelvic bones are not preserved but there are slight traces of the small pelvic fins under the cloacal region. Axial skeleton (Figs 1���2, 7) Starting from the tail region, the axial skeleton progressively elevates to reach anteriorly the level of the orbit. The vertebrae are formed by the dorsal and ventral arcocentra. There are no ossified vertebral centra, as in all pycnodonts. The neural and haemal arches are small and there is no dorso-ventral contact between them. So, the notochord is not completely surrounded by bony elements. In the caudal region, one small pre- and one small postzygapophysis link each neural arch with the following one. There are 25 neural spines before the epichordal series and 12 haemal spines before the hypochordal pieces. These spines bear anterior sagittal wings, except the fifth to the eighth neural spines that are pressed the ones against the others. The first 8 neural spines are autogenous. There are at least 11 pairs of long ribs. These ribs do not reach the ventral margin of the body. The postcoelomic bone is long and very narrow, except in its ventral part that is greatly enlarged. Dorsal and anal fins (Figs 1���2) Both the dorsal and the anal fins are long and strip-like (type A 2 of Poyato-Ariza& Wenz2002: fig. 34). The origin of the dorsal fin is located far from the head. The dorsal fin is supported by 42 pterygiophores and the anal fin by 34 pterygiophores. Only a few fragments of the dorsal and anal fins are preserved. The precise number of these rays is not known. Caudal skeleton (Figs 8���9) The caudal peduncle is very short, the dorsal and anal fins ending near the tail. The caudal endoskeleton is composed of 5 epichordals, 11 hypochordals and 1 urodermal. The first three epichordals are thin and elongate. The last two elements of the series are extremely reduced. The hypochordals are longer and wider than the epichordals. The sixth and the ninth hypochordals are moderately broadened. The caudal fin is double emarginated (Poyato-Ariza& Wenz2002: fig. 36E). There are 20 principal caudal rays, five dorsal and five ventral procurrent rays. Squamation (Figs 2, 10���11) Flank scales are present only in the abdominal region of the body. Dorsally, there is a pair of elongate scale bars associated to each dorsal ridge scute, except the last one. The squamation is missing in the middle region of the body. Ventrally, the flank scale bars are shorter but more numerous. The most ventral scales are small and triangular in shape. They are associated with the ventral keel scutes. A small scale, probably bearing the lateral line canal, is visible below the brush-like process of the parietal. There is a small narrow bifid cloacal scale and one broader postcloacal scale. The bifid cloacal scale is surrounded by a series of small scale bars. A long and rather broad scale is present just before the cloacal region. The dorsal ridge is formed by 11 scutes. The first dorsal scute is articulated with the dermosupraoccipital and is longer than the following scutes. The first two elements of the series bear a small spine. The ventral keel contains 13 scutes of which 11 prepelvic and two postcloacal associated to the postcoelomic bone. The 11 prepelvic scutes are very badly preserved and their shape is not determinable. The first postcloacal scute exhibits three small spines., Published as part of Taverne, Louis & Capasso, Luigi, 2018, Osteology and relationships of Libanopycnodus wenzi gen. et sp. nov. and Sigmapycnodus giganteus gen. et sp. nov. (Pycnodontiformes) from the Late Cretaceous of Lebanon, pp. 1-29 in European Journal of Taxonomy 420 on pages 4-14, DOI: 10.5852/ejt.2018.420, http://zenodo.org/record/1210447, {"references":["Taverne L. 1997. Les poissons cretaces de Nardo. 5 °. Pycnodus nardoensis sp. nov. et considerations sur l'osteologie du genre Pycnodus (Actinopterygii, Halecostomi, Pycnodontiformes). Bollettino del Museo Civico di Storia Naturale di Verona 21: 437 - 454.","Poyato-Ariza F. J. & Wenz S. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139 - 248."]}

Genus Libanopycnodus gen. nov. urn:lsid:zoobank.org:act:30B10E58-4147-4982-ADE0-B5956A502764 Type species Libanopycnodus wenzi gen. and sp. nov. (by monotypy). Diagnosis As for the species (monospecific genus). Etymology The generic eptithet refers to Lebanon and the taxon name Pycnodus is added.

Libanopycnodus wenzi gen. et sp. nov. urn:lsid:zoobank.org:act:2AD09A18-0416-4A26-AD08-E5307B901759 Figs 1–11 Diagnosis Small, deep-bodied pycnodontid fish, with rounded dorsal and ventral profiles. Head triangular in shape, with a rectilinear frontal profile and a short postorbital region. Pointed snout, with a mouth gape ventrally inclined. Frontal short. Prefrontal very narrow. Dorsal region of parietal forming a marked angle with the ventral region. Parietal brush-like process present. No temporal fenestra. Dermosphenotic sutured to the skull roof. Large, deep, well-visible dilatator fossa surrounded by dermosphenotic and dermopterotic. Posterior region of the endocranium exposed behind the dermopterotic. Vomer with 10 rounded molariform teeth on the external dental row. Premaxilla bearing two incisiform teeth. Prearticular with three rows of molariform teeth. Ectopterygoid present. Large triangular first infraorbital. Preopercle much deeper than the exposed region of the hyomandibula-dermohyomandibula. Opercle small and coma-shaped. Notochord not completely surrounded by vertebral arches. Neural and haemal spines with an anterior wing-like component. Twenty-five neural spines before epichordal series. First eight neural spines autogenous. Twelve haemal spines before hypochordal series. Eleven pairs of ribs. Postcoelomic bone long, narrow, with an enlarged ventral extremity. Dorsal and anal fins strip-like. Dorsal fin supported by 42 pterygiophores. Anal fin supported by 34 pterygiophores. Caudal peduncle short. Five epichordals. Eleven hypochordals, with two moderately broadened. One urodermal. Caudal fin double emarginated, with 20 principal rays. A few scale bars on the abdominal region of the body. Small complete scales associated with the prepelvic ventral keel scutes. Bifid cloacal scale present. One postcloacal scale. Dorsal ridge with 11 scutes, of which the first two with a small spine. Ventral keel with 13 scutes, 11 prepelvic and two postcloacal with small spines. Etymology The specific epithet is in homage to Dr. Sylvie Wenz (Paris), who worked abundantly on the pycnodontiform fishes. Material examined Holotype LEBANON: a complete specimen (Figs 1–2), total length: 90 mm, standard length: 78 mm, marine Upper Cenomanian deposits of Ein Namoura (CLC S-574). General morphology and morphometric data The fish is deep-bodied, the maximum body height being equal to a little more than the half of the standard length. The dorsal and the ventral profiles are rounded and devoid of marked apex. The morphometric data are given in % of the standard length (78 mm): Length of the head (opercle included) ……………………………………………………………34.0 % Depth of the head (in the occipital region) ………………………………………………………44.7 % Maximum depth of the body (just behind the head) ……………………………………………56.2 % Prepelvic length …………………………………………………………………………………57.0 % Predorsal length …………………………………………………………………………………68.1 % Basal length of the dorsal fin ……………………………………………………………………35.7 % Preanal length ……………………………………………………………………………………71.5 % Basal length of the anal fin ………………………………………………………………………28.9 % Depth of the caudal peduncle ………………………………………………………………………7.2 % Osteology Skull (Figs 3–6) The head is large, triangular in shape, altogether long and deep, with an almost rectilinear frontal profile. The preorbital region is much longer than the postorbital one. The orbit is wide. The dermal bones of the skull are ornamented with small tubercles. The snout is pointed, with a mouth gape ventrally inclined. The mesethmoid is elongate and broad. Its upper margin is covered posteriorly by a pair of very narrow prefrontals and anteriorly by the premaxillae. The orbitosphenoid is pressed against the posterior margin of the mesethmoid. The vomer, seen in profile, is long and deeper posteriorly than anteriorly. Only the lateral dental row, composed of 7 well-developed, rounded molariform teeth, is visible. There are traces of 3 other very small teeth located at the anterior extremity of the lower margin of the vomer. The frontal is short and limited to the orbital region. The dermosupraoccipital is small, with a pointed dorso-posterior extremity. The parietal exhibits an unusual and strange shape. Its dorsal half forms a marked angle with its ventral part. This peculiar shape of the parietal is not due to an artefact of fossilisation. A short brush-like process (= branched peniculus) is attached to the parietal. The supratemporal is a rather broad scale that lies against the upper part of the parietal, just below the first dorsal ridge scute. The dermopterotic is deeper than long. The dermosphenotic is not a free bone. It is included in the lateral wall of the skull roof and is sutured with the frontal and the dermopterotic. The ventral margin of the dermosphenotic and the dermopterotic are located at the same level as the lower border of the orbit. The dermopterotic and the dermosphenotic surround a deep and wide dilatator fossa. The parasphenoid is long, broad, toothless and its trabercular region is obliquely inclined. The posterior region of the endocranium is well visible. A small part of the intercalar and the exoccipital are preserved behind the lower half of the parietal. The ventral margin of the exoccipital is pierced by a large foramen for the branches of the vagus nerve (X). The basioccipital, articulated with the first basiventral, is also present. The exoccipital and basioccipital are separated from the braincase main part as in Pseudopycnodus nardoensis (Taverne 1997: fig. 2). A cartilaginous region probably linked the two cranial regions on the living fish. The autosphenotic and the sensory canals of the skull roof are not visible. A temporal fenestra is not present. The quadratic arch includes wide metapterygoid and entopterygoid plus a small ectopterygoid. Only the anterior parts of the quadrate and the symplectic are preserved. Both bones articulate with the lower jaw. The articular head of the symplectic is very broad. The premaxilla is long and narrow. It bears two incisiform teeth. The maxilla is lost. The posterior extremity of the dentary is preserved but not its anterior part with the teeth. The articular is small. The angular is not preserved. The prearticular is a large bone with a marked hook-like coronoid process, bearing three rows of molariform teeth. The dorsal row contains 6 teeth, the middle row 7 teeth and the ventral row at least 5 teeth. The teeth of the dorsal row are the smallest, with a slightly concave surface and their dorsal margin bears some little tubercles. The teeth of the lower row are the largest. They are deeper than long, with a slightly convex surface. The orbital series contains four bones. The first infraorbital is a large triangular bone. The second and third infraorbitals are tubular bones. As already written, the dermosphenotic is sutured to the skull roof. A few fragments of the sclerotic bony ring are visible just under the frontal. The exposed part of the hyomandibula-dermohyomandibula is much smaller than the preopercle to which it is sutured. The long ventral branch of the hyomandibula is covered by the preopercle but remains clearly visible. The upper margin of the hyomandibula bears a broad dorsal process that articulates in the dilatator fossa. The preopercle is deep and broad, with a rather narrow dorsal margin. The opercle is well developed, with an acuminate ventral tip and a broader upper part. The anterior ceratohyal and two branchiostegal rays are visible below the preopercle. Girdles (Figs 3–4) A small posttemporal is preserved behind the skull, at the level of the suture between the parietal and the dermopterotic. The hypercleithrum (= supracleithrum) is a long and thin bone. The cleithrum is divided in a short but very broad ventral branch and an extremely long but narrow dorsal branch. No pterygiophore is preserved but fragments of short pectoral rays are visible. The pelvic bones are not preserved but there are slight traces of the small pelvic fins under the cloacal region. Axial skeleton (Figs 1–2, 7) Starting from the tail region, the axial skeleton progressively elevates to reach anteriorly the level of the orbit. The vertebrae are formed by the dorsal and ventral arcocentra. There are no ossified vertebral centra, as in all pycnodonts. The neural and haemal arches are small and there is no dorso-ventral contact between them. So, the notochord is not completely surrounded by bony elements. In the caudal region, one small pre- and one small postzygapophysis link each neural arch with the following one. There are 25 neural spines before the epichordal series and 12 haemal spines before the hypochordal pieces. These spines bear anterior sagittal wings, except the fifth to the eighth neural spines that are pressed the ones against the others. The first 8 neural spines are autogenous. There are at least 11 pairs of long ribs. These ribs do not reach the ventral margin of the body. The postcoelomic bone is long and very narrow, except in its ventral part that is greatly enlarged. Dorsal and anal fins (Figs 1–2) Both the dorsal and the anal fins are long and strip-like (type A 2 of Poyato-Ariza& Wenz2002: fig. 34). The origin of the dorsal fin is located far from the head. The dorsal fin is supported by 42 pterygiophores and the anal fin by 34 pterygiophores. Only a few fragments of the dorsal and anal fins are preserved. The precise number of these rays is not known. Caudal skeleton (Figs 8–9) The caudal peduncle is very short, the dorsal and anal fins ending near the tail. The caudal endoskeleton is composed of 5 epichordals, 11 hypochordals and 1 urodermal. The first three epichordals are thin and elongate. The last two elements of the series are extremely reduced. The hypochordals are longer and wider than the epichordals. The sixth and the ninth hypochordals are moderately broadened. The caudal fin is double emarginated (Poyato-Ariza& Wenz2002: fig. 36E). There are 20 principal caudal rays, five dorsal and five ventral procurrent rays. Squamation (Figs 2, 10–11) Flank scales are present only in the abdominal region of the body. Dorsally, there is a pair of elongate scale bars associated to each dorsal ridge scute, except the last one. The squamation is missing in the middle region of the body. Ventrally, the flank scale bars are shorter but more numerous. The most ventral scales are small and triangular in shape. They are associated with the ventral keel scutes. A small scale, probably bearing the lateral line canal, is visible below the brush-like process of the parietal. There is a small narrow bifid cloacal scale and one broader postcloacal scale. The bifid cloacal scale is surrounded by a series of small scale bars. A long and rather broad scale is present just before the cloacal region. The dorsal ridge is formed by 11 scutes. The first dorsal scute is articulated with the dermosupraoccipital and is longer than the following scutes. The first two elements of the series bear a small spine. The ventral keel contains 13 scutes of which 11 prepelvic and two postcloacal associated to the postcoelomic bone. The 11 prepelvic scutes are very badly preserved and their shape is not determinable. The first postcloacal scute exhibits three small spines.

Taverne, Louis, Capasso, Luigi (2018): Osteology and relationships of Libanopycnodus wenzi gen. et sp. nov. and Sigmapycnodus giganteus gen. et sp. nov. (Pycnodontiformes) from the Late Cretaceous of Lebanon. European Journal of Taxonomy 420: 1-29, DOI: https://doi.org/10.5852/ejt.2018.420

Genus Sigmapycnodus gen. nov. urn:lsid:zoobank.org:act:6855D3A7-D247-4065-81D2-26D5BD0ACF63 Type species Sigmapycnodus giganteus gen. and sp. nov. (by monotypy). Diagnosis As for the species (monospecific genus). Etymology From the Greek letter ‘ sigma ’. The taxon name Pycnodus is added. The generic name refers to the sigmoid frontal profile of the new fossil fish.

Genus Sigmapycnodus gen. nov. urn:lsid:zoobank.org:act:6855D3A7-D247-4065-81D2-26D5BD0ACF63 Type species Sigmapycnodus giganteus gen. and sp. nov. (by monotypy). Diagnosis As for the species (monospecific genus). Etymology From the Greek letter ��� sigma ���. The taxon name Pycnodus is added. The generic name refers to the sigmoid frontal profile of the new fossil fish., Published as part of Taverne, Louis & Capasso, Luigi, 2018, Osteology and relationships of Libanopycnodus wenzi gen. et sp. nov. and Sigmapycnodus giganteus gen. et sp. nov. (Pycnodontiformes) from the Late Cretaceous of Lebanon, pp. 1-29 in European Journal of Taxonomy 420 on page 15, DOI: 10.5852/ejt.2018.420, http://zenodo.org/record/1210447

We hereafter use the phylogeny of Pycnodontiformes proposed by Poyato-Ariza & Wenz (2002, 2005) and based on cranial and postcranial characters. Brembodidae represents the most basal family and the most ancient lineage in the order (Figs 20–23; Tintori 1980; Nursall 1996, 1999; Poyato-Ariza & Wenz 2002). They date from the Upper Norian (Late Triassic) of North Italy. Two genera are known, Brembodus Tintori, 1980 and Gibbodon Tintori, 1980. They are deep-bodied fishes, with an important dorsal gibbosity or an elongate, spiny dorsal process. Brembodus still possesses two well developed dermosupraoccipitals, the posterior one being especially elongated. It is to be noted that Tintori (1980: fig. 1) considers the posterior dermosupraoccipital of Brembodus as the first scale of the dorsal ridge, and Nursall (1996: 145, character 94) and Poyato-Ariza & Wenz (2002: 195, character 84[1]) as a dorsal spine. However, this large bone not only articulates with, but is also sutured to the anterior dermosupraoccipital and the parietal, exactly as the posterior dermosupraoccipital in the five gyrodontiform genera. So, if this bone is considered as a posterior dermosupraoccipital in Gyrodontiformes, there is no valid reason to give it another name in B. ridens. The case of Gibbodon is more problematic. Tintori (1980: fig. 2) shows two dermosupraoccipitals in this genus but Poyato-Ariza & Wenz (2002, fig. 6 B) figure only one occipital bone preceding the first small dorsal scute. Both brembodid genera possess tubular posterior infraorbitals and a gigantic first infraorbital completely covering the cheek. They preserve small bony tesserae in the gular region. They have 3 teeth in the upper jaw and 4 or 5 teeth on the dentary. The margins of their unpaired fins bear fringing fulcra. There is a series of urodermals in the caudal skeleton. The scales cover their body totally as in Gyrodontiformes. However, these scales are much deeper than broad, with an important development of the bar component, while the flank scales of Gyrodus are less deep and the bar component is less marked (Hennig 1906: pl. 11;, Lambers 1991: fig. 3a). There is a mosaic of small scales in the cloacal region of Brembodidae as in Gyrodus (Poyato-Ariza & Wenz 2002: fig. 40A). The new Lebanese fossil fish and the other Pycnodontiformes share at least two apomorphic characters not present in Brembodidae, i.e., the loss of the fringing fulcra and the presence of scales only in the abdominal region of the body. Macromesodon Blake, 1905 (= Eomesodon Woodward, 1918 pro parte) seems to be the most basal member of this remaining group (Woodward 1918; Poyato-Ariza & Wenz 2002, 2004). There is a large dorsal prominence as in Brembodidae. Tubular infraorbitals are present, but some bony tesserae are still covering part of the cheek. There is a series of 5 or 6 urodermals in the caudal skeleton. The cloacal region is still covered by a mosaic of small scales as in the most plesiomorphic Pycnodontomorpha. All body scales are completely ossified. Macromesodon is generally devoid of fringing fulcra on the impaired fins. However some rare samples still exhibit a few fringing fulcra (Lambers 1991: fig. 25c; Poyato-Ariza & Wenz 2002: 192). Acrorhinichthys gen. nov. and the more advanced Pycnodontiformes exhibit some new apomorphic characters. A dermohyomandibula is fused with the hyomandibula. There are anterior sagittal flanges on the neural and haemal spines. At least some neural and haemal arches are connected to each other by means of one or more postzygapophyses. The ventral keel contains less than 18 scutes. The urodermal series is reduced to 2, 1 or 0 elements. A part of the body squamation is composed of scale bars. The number of scales in the cloacal region is greatly reduced. Acrorhinichthys gen. nov. is the most plesiomorphic genus within those remaining Pycnodontiformes. It still possesses a small dorsal prominence and a few gular bony tesserae, two plesiomorphic characters already disappeared in Coccodontidae, Gladiopycnodontidae, Gebrayelichthyidae and Pycnodontidae. Coccodontidae, Gladiopycnodontidae and Gebrayelichthyidae are three highly specialized families of Pycnodontiformes (Gayet 1984; Nursall & Capasso 2008; Capasso et al. 2010; Taverne & Capasso 2013a, 2014a, b, c). Their morphology and osteology are completely different from those of Acrorhinichthys gen. nov. Pycnodontidae are principally characterized by the brush-like process of the parietal, a structure absent in Acrorhinichthys gen. nov. However, Akromystax Poyato-Ariza & Wenz, 2005 from the Cenomanian of Lebanon (Fig. 24), the most plesiomorphic genus within Pycnodontidae, has preserved some archaic characters absent in the more evolved Pycnodontidae but present in Acrorhinichthys gen. nov., such as a series of complete scales associated with the dorsal ridge scutes (Poyato-Ariza & Wenz 2005: fig. 2), two imbricated complete cloacal scales, a small ventral triangular one and a deeper dorsal, one with a well developed bar component and a broad concave lower margin (Fig. 25), and more than two teeth on the premaxilla and the dentary (Poyato-Ariza & Wenz 2005: fig. 7).

Joinvillichthys kriweti sp. nov. urn:lsid:zoobank.org:act: B8C4822F-44CA-4776-8455-452C5DBDCA1C Figs 12-16 Diagnosis Joinvillichthys with a body depth equal to 46.0 % of the standard length. A dorsal prominence present on the frontal. Maxilla elongated. Large parietal. Dermosupraoccipital sutured to the parietal and not to the dermopterotic. Dermopterotic much longer than deep. Small dermosphenotic. Comma-shaped opercle. Anterior ventral branch of the cleithrum lost. Broad and short pectoral spine articulated on the ventral margin of the cleithral posterior process. Caudal fin double emarginated. Etymology The name of the new species is dedicated to Dr Jürgen Kriwet (Vienna) who has greatly improved our knowledge of the pycnodontiform fishes. Holotype LEBANON: sample CLC S-137, a complete specimen from Haqel (Fig. 12), total length: 91 mm. Paratype LEBANON: sample AMNH 4517a (3698) and counterpart, an incomplete specimen from Hgula (Hay 1903: pl. 29, fig. 1); only the head and the beginning of the body are preserved, length: 63 mm. Formation and locality Marine Upper Cenomanian, Haqel and Hgula, Lebanon. Morphometric data (Fig. 13) The morphometric data are given in % of the holotype standard length (76 mm) Length of the head (dermosupraoccipital included) …………………………………………… 54.2 % Length of the cephalo-thorax (cleithrum included) ………………………………………… 75.3 % Depth of the head (without the nuchal horn) ……………………………………………… 43.8 % Length of the nuchal horn …………………………………………………………………… 27.7 % Maximum depth of the body (just behind the nuchal horn) ………………………………… 46.0 % Predorsal length ………………………………………………………………………………… 76.6 % Basal length of the dorsal fin …………………………………………………………………… 9.8 % Preanal length …………………………………………………………………………………… 80.4 % Basal length of the anal fin ………………………………………………………………………… 6.4 % Depth of the caudal peduncle ……………………………………………………………………… 9.8 % Osteology 1. The skull (Fig. 14) The general morphology of the skull is rather close to that of Joinvillichthys lindstroemi and the cranial dermal bones also are ornamented with small tubercles. However, there are many small differences in the head skeleton of the two fishes. Thus the description that follows will principally emphasize these differences. The skull is shorter and deeper than in Joinvillichthys lindstroemi. Its depth, from the upper margin of the dermosupraoccipital to the lower margin of the cleithrum, is equal to 83 to 86 % of its length, from the tip of the snout to the basis of the nuchal horn. The rostrum lengthening is less pronounced. The prefrontal is broader and has a very sinuous suture with the frontal. Its anterior tip also bears very small spines but is less outpacing of the lower jaw level. The frontal is broader but does not outpace posteriorly the level of the orbit. The bone bears a small dorsal prominence. The dermosupraoccipital is longer and is sutured to the parietal and the supratemporal but not with the dermopterotic. The parietal is considerably larger. The dermopterotic is longer but much thinner. The supratemporal is sutured to the parietal and reaches the dermopterotic at only one point. As in Joinvillichthys lindstroemi, the long nuchal horn is supported only by the dermosupraoccipital. The orbitosphenoid and the pleurosphenoid are present in the orbit, just below the frontal, but the basisphenoid is not visible. The toothless premaxilla is longer and narrower. The toothless maxilla also is narrower and more elongate. The lower jaw is composed with the same bones but is longer. The dentary bears two small incisiform teeth and its ventral margin is denticulated. The articulation with the quadrate is located at the level of the posterior border of the orbit. A fragment of a large first infraorbital is preserved on the suture between the prefrontal and the premaxilla. The sclerotic ring is visible in the orbit. The hyomandibula and the preopercle are sutured together. The exposed part of the hyomandibuladermohyomandibula is larger than in Joinvillichthys lindstroemi but still much smaller than the considerably enlarged preopercle. The opercle is broader and comma-shaped, with the sharp end dorsally located. A part of the anterior ceratohyal and two small branchiostegal rays are visible behind the lower jaw. 2. The girdles (Figs 14, 16) The bones of the gigantic pectoral girdle have the same size and shape as in Joinvillichthys lindstroemi. However, two important differences exist. The anterior ventral branch of the cleithrum is lost. No postcleithrum is visible, but that is perhaps due to the fossilization. The pectoral spine is shorter, much broader and is not articulated with the rear of the cleithrum but more anteriorly on its lower margin. A small pelvic girdle is present. Indeed, a part of a vertically oriented pelvic bone is visible under a broken part of the cleithrum. 3. The axial skeleton (Fig. 13) The trunk is fusiform but proportionally deeper than in Joinvillichthys lindstroemi. The axial skeleton is incomplete. Three vertebral segments are missing near the caudal peduncle. There are 16 neural spines (the three missing ones included) before the epichordal series. Only 8 haemal spines are preserved. The total number of haemal spines must be around 12 or 13. The neural and haemal spines are short but broad. The neural and haemal arches surround almost completely the notochord. No ribs are visible. The postcoelomic bone is well developed and backwardly oriented. 4. The dorsal and anal fins (Fig. 13) The short dorsal and anal fins are located at the mid-length of the body. There are 9 pterygiophores and 9 rays in the dorsal fin. The anal fin contains 7 rays, but the number of anal pterygiophores is not determinable. The first dorsal and anal ray is spiny. The other rays are segmented. 5. The caudal skeleton (Fig. 15) The caudal skeleton of the holotype is partly preserved. There are 6 short and broad epichordals and 7 hypochordals. However, one or two anterior hypochordals are missing, so the complete series must be composed of 8 or 9 elements. The fifth preserved hypochordal is strongly enlarged. No urodermal is visible, but the region where theses bones are usually present is not preserved. The caudal fin is double emarginated (Poyato-Ariza & Wenz 2002: fig. 36E) and contains 19 principal segmented caudal rays, the 2 external being pointed and the 17 others branched. There are 6 ventral and at least 4 dorsal procurrent rays. 6. Squamation (Fig. 16) The squamation is the same as in Joinvillichthys lindstroemi. There are 7 spiny scutes in the dorsal ridge and at least 3 spiny scutes in the ventral keel. The two posterior ventral scutes are associated with the ventral margin of the postcoelomic bone. The body scales are slightly ornamented with tubercles. Anteriorly, they are small and flake-like. Posteriorly, there are much larger, irregular and scute-like shaped.

Joinvillichthys gen. nov. urn:lsid:zoobank.org:act: 6D820D8E-DF5C-41E9-B351-F46F7A9D4119 Type species: Coccodus lindstroemi Davis, 1890 (here designated). Diagnosis Gladiopycnodontid with an elongate prefrontal forming a short rostrum outpacing the lower jaw level. Anterior extremity of the prefrontal acuminate and spiny. Vomer bearing small rounded molariform teeth irregularly ranged. Orbitosphenoid, pleurosphenoid and basisphenoid present. Supratemporal sutured to the rear of the skull. Premaxilla long, broad, toothless and sutured by its upper margin to the prefrontal. Dentary bearing 2 incisiform teeth. Hypertrophied trapezoid preopercle covering the cheek. Large first infraorbital (only known in Joinvillichthys kriweti gen. et sp. nov.). Exposed part of the hyomandibula-dermohyomandibula much smaller than the preopercle. Long nuchal horn with a spiny posterior margin and articulated only on the dermosupraoccipital. Pectoral girdle closely associated to the skull, forming a cephalo-thorax. Cleithrum hypertrophied, with a gigantic posterior ventral process. Hypercleithrum hypertrophied. Well developed posttemporal, with an acuminate posterior extremity. 16 to 17 neural spines, all fused to the neural arches, before the epichordal series. 10 haemal spines before the hypochordal series (only known in Joinvillichthys lindstroemi). Short dorsal fin with 8 or 9 rays. Short anal fin with 7 to 9 rays. 7 to 9 spiny scutes in the dorsal ridge. 3 or 4 scutes in the ventral keel, the last scute or the two last ones associated to the postcoelomic bone. Body completely covered by small, flake-like scales in the abdominal region and by large, scute-like scales in the caudal region. Etymology The generic name is chosen in memory of Lord Jean de Joinville (1224–1317), seneschal of Champagne, who related in his biography of Louis IX the presentation of some Lebanese fossil fishes to this French king at Saïda during the seventh crusade (Gayet et al. 2012: 8). The Greek word ichthys, fish, is added., Published as part of Taverne, Louis & Capasso, Luigi, 2014, On the " Coccodus " lindstroemi species complex (Pycnodontiformes, Gladiopycnodontidae) from the marine Late Cretaceous of Lebanon, with the description of two new genera, pp. 1-27 in European Journal of Taxonomy 101 on page 4, DOI: 10.5852/ejt.2014.101, http://zenodo.org/record/3838770, {"references":["Davis J. W. 1890. On a new species of Coccodus (C. lindstroemi, Davis). The Quarterly Journal of the Geological Society of London 46 (34): 565 - 568.","Gayet M., Abi Saad P. & Gaudant O. 2012. Les fossiles du Liban. Memoire du temps. Editions Desiris, Meolan-Revel."]}

Joinvillichthys lindstroemi (Davis, 1890) Figs 1-11 Diagnosis Joinvillichthys with a body depth comprising between 23.8 and 34.0 % of the standard length. No dorsal prominence on the frontal. Maxilla triangular in shape. Small parietal. Dermosupraoccipital sutured with the parietal and the dermopterotic. Dermopterotic deeper than long. Large dermosphenotic. Thin, rodlike opercle. Anterior ventral branch of the cleithrum present. Thin pectoral spine articulated on the rear of the cleithral posterior process. Caudal fin with a convex posterior margin. Synonymy Coccodus Lindstroemi Davis, 1890: 567, pl. 22. ��� Coccodus ��� lindstroemi ��� Poyato-Ariza & Wenz, 2002: 145. Holotype LEBANON: sample NRM PZ P. 2073, a complete specimen from Haqel (Fig. 1), total length: 76 mm. Other specimens LEBANON: sample IRSNB N�� P 9276, a nearly complete specimen (the caudal fin is missing) from Hgula (Fig. 2), length: 93 mm; sample CLC S-138, a nearly complete specimen (a part of the caudal fin is missing) from Haqel (Fig. 3), length: 82 mm; sample CLC S-324, a complete specimen from Haqel, total length: 76 mm. Formation and locality Marine Upper Cenomanian, Haqel and Hgula, Lebanon. Morphometric data (Fig. 4) The morphometric data are given in % of the standard length for the holotype NRM PZ P. 2073 (64 mm) and for sample IRSNB N�� P 9276 (93 mm). These two specimens represent the two extremes of the species morphometric variation, as measured on the four studied samples Holotype P9276 Length of the head (dermosupraoccipital included)............................................. 55.8 % 56.3 % Length of the cephalo-thorax (cleithrum included).............................................. 63.9 % 68.1 % Depth of the head (without the nuchal horn)........................................................ 35.4 % 25.0 % Length of the nuchal horn..................................................................................... 25.2 % 25.6 % Maximum depth of the body (just behind the nuchal horn)................................. 34.0 % 23.8 % Predorsal length.................................................................................................... 74.8 % 73.1 % Basal length of the dorsal fin................................................................................ 9.5 % 10.6 % Preanal length....................................................................................................... 72.1 % 73.7 % Basal length of the anal fin................................................................................... 8.2 % 9.4 % Depth of the caudal peduncle............................................................................... 5.4 % 5.3 % The important individual differences in the values of the head and body depths are probably due to sexual or seasonal variations. Osteology 1. The skull (Figs 5���8) The head is very large. Its length, from the tip of the snout to the basis of the nuchal horn, is equal to the body length. According to the specimens, its depth, from the upper margin of the dermosupraoccipital to the lower margin of the cleithrum, represents from 48 to 67 % of its length. The dermal bones of the skull are ornamented with small tubercles. The long, pointed rostrum slightly outpaces the lower jaw and is formed by two large paired bones, the prefrontal and the premaxilla. The anterior tip of the prefrontal bears two or three very small spines. Posteriorly, the bone reaches the orbit level. Only the most posterior part of the mesethmoid is visible. The vomer is completely hidden by the premaxilla. However, a small anterior fragment of the premaxilla is lost on sample CLC S-138 and a part of the vomer is visible. The bone bears small, rounded molariform teeth that are irregularly ranged. The frontal is short, not curved and slightly broader posteriorly than anteriorly. The posterior portion of the frontal outpaces the level of the orbit. The posterior lateral part of the skull roof is formed on each side by four small bones, the parietal, the dermosphenotic, the dermopterotic and the supratemporal. The dermosphenotic partly covers the autosphenotic. The dermopterotic is deeper than long. The supratemporal is sutured to the dermopterotic and does not reach the parietal. The dermosupraoccipital occupies the median posterior part of the skull roof.This large bone is sutured with the frontal, the parietal, the dermopterotic and the supratemporal. A long pointed nuchal horn is fixed to the dermosupraoccipital. This horn is ornamented with long and thin striations and bears a series of spines on its posterior border. Sample IRSNB N�� P 9276 clearly shows the orbitosphenoid, the pleurosphenoid and the basisphenoid in the orbit. The three bones are pressed against the frontal. The orbitosphenoid reaches anteriorly the mesethmoid. The small basisphenoid is divided in a dorsal meningost and a short ventral belophragm. Fig. . The parasphenoid is very long, almost straight, but it does not reach the posterior border of the skull that is occupied by the basioccipital, as seen on the same specimen. Sample P 9276 also shows the very small prootic with a large foramen for the trigeminal nerve (V) in its anterior border. The anterior margin of the metapterygoid and the entopterygoid is visible between the preopercle and the parasphenoid. The quadrate and the symplectic remain hidden by the preopercle and the cleithrum. The premaxilla and the maxilla compose the upper jaw. As in other pycnodontomorph fishes, there is no supramaxilla. The broad, long and toothless premaxilla is located below the prefrontal to which it is sutured by its upper margin. The maxilla is large, toothless and triangle-shaped. The lower jaw comprises the dentary, the prearticular, the angular and the articular. The articulation with the quadrate is located at the level of the anterior border of the orbit. The prearticular is the largest bone of the series but is partly covered by the maxilla and the preopercle. The teeth of the prearticular are not visible. The articular and the angular are small bones. The dentary bears two incisiform teeth and is reduced to its ventral branch. Its lower margin is denticulated. The orbit is large and longer than deep. No orbital bone is preserved, except the dermosphenotic that is sutured to the frontal, the parietal and the dermopterotic. Fragments of a sclerotic bony ring are visible on the holotype. The hyomandibula-dermohyomandibula and the preopercle are sutured together. The exposed part of the hyomandibula-dermohyomandibula is much smaller than the greatly enlarged preopercle. The opercle is a long and very thin bone wedged between the preopercle and the cleithrum. Small fragments of branchial bones with a few long and acuminate branchiospines are visible on sample IRSNB N�� P 9276. 2. The girdles (Figs 4���6) As in all Gladiopycnodontidae, the enlarged pectoral girdle is closely associated to the skull, forming a sort of cephalo-thorax. The dermal bones are ornamented with small tubercles. The postemporal rests on the large ovoid hypercleithrum (= supracleithrum) and is articulated with the dermosupraoccipital by its broad anterior border. Its posterior extremity is acuminate. The cleithrum is enormous, with a well developed anterior branch and a very wide posterior process. There is a small postcleithrum. The pectoral fin is replaced by a long and thin spine that is decorated with a few ridges and tubercles. The spine is articulated on the rear of the cleithral posterior process. No trace of a pelvic girdle is visible. It is possible that reduced pelvic bones and fins were present but hidden by the gigantic cleithral posterior process. Such a situation exists in other gladiopycnodontid fishes (Taverne & Capasso 2013: figs 8, 18). 3. The axial skeleton (Fig. 4) As in most Gladiopycnodontidae, the trunk is fusiform and not deep-bodied. Sample IRSNB N�� P 9276 has lost the scales on the body and so the well preserved axial skeleton is completely accessible. The vertebrae are constituted by the dorsal and ventral arcocentra. They surround almost completely the notochord. There are 17 neural spines before the epichordal elements and 10 haemal spines before the hypochordal series. Before the level of the dorsal fin, the neural spines are long and narrow. Posteriorly, the neural spines are much shorter but also a little broader. The haemal spines are short and broad. The number of ribs is not determinable but ribs are present under the cleithral posterior process, as seen on sample CLC S-138. The last ribs are very short. The postcoelomic bone is backwardly oriented and is articulated with the ventral arcocentrum preceding the one bearing the first haemal spine. The bone is broader ventrally than dorsally. 4. The dorsal and anal fins (Fig. 4) The dorsal and anal fins are short and located in the middle of the body length. There are 8 or 9 dorsal pterygiophores and also 8 or 9 anal pterygiophores, each of them bearing a ray. The first dorsal and anal ray is spiny. The other rays are segmented. 5. The caudal skeleton (Figs 9���10) Sample IRSNB N�� P 9276 presents the best preserved caudal skeleton. The caudal peduncle is long and includes 5 or 6 vertebral segments. There are 6 epichordals and a least 8 hypochordals. The hypochordals are broader than the long, thin and pointed epichordals. In specimen IRSNB N�� P 9276, the sixth and seventh hypochordals are moderately broadened and partly fused together. In sample CLC S-138, the sixth and seventh hypochordals are not fused and the broadening only exists on the seventh element. The caudal fin has a convex posterior margin (Poyato-Ariza & Wenz 2002: fig. 36B) and contains 17 or 18 principal caudal rays. There are a few procurrent rays in each lobe. 6. Squamation (Fig. 11) The body is entirely covered by scales ornamented with tubercles and imbricated one into another. Anteriorly, these scales are very small, flake-like and they extend on the cleithral posterior process. Posteriorly to the median fins, these scales are a much larger, irregular and scute-like shaped. Between the nuchal horn and the dorsal fin, the dorsal ridge is composed by 7 to 9 scutes with a spiny upper margin. The ventral keel contains at least 4 scutes. The first three are located in the cloacal region. The posterior one of these three has a spiny lower margin. A fourth spiny scute is associated with the ventral extremity of the postcoelomic bone., Published as part of Taverne, Louis & Capasso, Luigi, 2014, On the " Coccodus " lindstroemi species complex (Pycnodontiformes, Gladiopycnodontidae) from the marine Late Cretaceous of Lebanon, with the description of two new genera, pp. 1-27 in European Journal of Taxonomy 101 on pages 4-13, DOI: 10.5852/ejt.2014.101, http://zenodo.org/record/3838770, {"references":["Davis J. W. 1890. On a new species of Coccodus (C. lindstroemi, Davis). The Quarterly Journal of the Geological Society of London 46 (34): 565 - 568.","Poyato-Ariza F. J. & Wenz S. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139 - 248.","Taverne L. & Capasso L. 2013. Gladiopycnodontidae, a new family of pycnodontiform fishes from the Late Cretaceous of Lebanon, with the description of three genera. European Journal of Taxonomy 57: 1 - 30. http: // dx. doi. org / 10.5852 / ejt. 2013.57"]}

Pankowskichthys gen. nov. urn:lsid:zoobank.org:act: 6C6C231C-40DC-4AD6-9250-2FD72E4EDC1B Type species: Pankowskichthys libanicus gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology The name of the new genus is dedicated to Mark Pankowski (Rockville, Maryland, U.S.A.), who generously offered the holotype of Pankowskichthys libanicus gen. et sp. nov. to the Royal Belgian Institute for Natural Sciences (IRSNB)., Published as part of Taverne, Louis & Capasso, Luigi, 2014, On the " Coccodus " lindstroemi species complex (Pycnodontiformes, Gladiopycnodontidae) from the marine Late Cretaceous of Lebanon, with the description of two new genera, pp. 1-27 in European Journal of Taxonomy 101 on page 19, DOI: 10.5852/ejt.2014.101, http://zenodo.org/record/3838770

Pankowskichthys gen. nov. urn:lsid:zoobank.org:act: 6C6C231C-40DC-4AD6-9250-2FD72E4EDC1B Type species: Pankowskichthys libanicus gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology The name of the new genus is dedicated to Mark Pankowski (Rockville, Maryland, U.S.A.), who generously offered the holotype of Pankowskichthys libanicus gen. et sp. nov. to the Royal Belgian Institute for Natural Sciences (IRSNB).

Joinvillichthys kriweti sp. nov. urn:lsid:zoobank.org:act: B8C4822F-44CA-4776-8455-452C5DBDCA1C Figs 12-16 Diagnosis Joinvillichthys with a body depth equal to 46.0 % of the standard length. A dorsal prominence present on the frontal. Maxilla elongated. Large parietal. Dermosupraoccipital sutured to the parietal and not to the dermopterotic. Dermopterotic much longer than deep. Small dermosphenotic. Comma-shaped opercle. Anterior ventral branch of the cleithrum lost. Broad and short pectoral spine articulated on the ventral margin of the cleithral posterior process. Caudal fin double emarginated. Etymology The name of the new species is dedicated to Dr J��rgen Kriwet (Vienna) who has greatly improved our knowledge of the pycnodontiform fishes. Holotype LEBANON: sample CLC S-137, a complete specimen from Haqel (Fig. 12), total length: 91 mm. Paratype LEBANON: sample AMNH 4517a (3698) and counterpart, an incomplete specimen from Hgula (Hay 1903: pl. 29, fig. 1); only the head and the beginning of the body are preserved, length: 63 mm. Formation and locality Marine Upper Cenomanian, Haqel and Hgula, Lebanon. Morphometric data (Fig. 13) The morphometric data are given in % of the holotype standard length (76 mm) Length of the head (dermosupraoccipital included) ��������������������������������������������������� 54.2 % Length of the cephalo-thorax (cleithrum included) ������������������������������������������������ 75.3 % Depth of the head (without the nuchal horn) ������������������������������������������������������ 43.8 % Length of the nuchal horn ������������������������������������������������������������������������������ 27.7 % Maximum depth of the body (just behind the nuchal horn) ��������������������������������������� 46.0 % Predorsal length ��������������������������������������������������������������������������������������������� 76.6 % Basal length of the dorsal fin ������������������������������������������������������������������������������ 9.8 % Preanal length ������������������������������������������������������������������������������������������������ 80.4 % Basal length of the anal fin ������������������������������������������������������������������������������������ 6.4 % Depth of the caudal peduncle ��������������������������������������������������������������������������������� 9.8 % Osteology 1. The skull (Fig. 14) The general morphology of the skull is rather close to that of Joinvillichthys lindstroemi and the cranial dermal bones also are ornamented with small tubercles. However, there are many small differences in the head skeleton of the two fishes. Thus the description that follows will principally emphasize these differences. The skull is shorter and deeper than in Joinvillichthys lindstroemi. Its depth, from the upper margin of the dermosupraoccipital to the lower margin of the cleithrum, is equal to 83 to 86 % of its length, from the tip of the snout to the basis of the nuchal horn. The rostrum lengthening is less pronounced. The prefrontal is broader and has a very sinuous suture with the frontal. Its anterior tip also bears very small spines but is less outpacing of the lower jaw level. The frontal is broader but does not outpace posteriorly the level of the orbit. The bone bears a small dorsal prominence. The dermosupraoccipital is longer and is sutured to the parietal and the supratemporal but not with the dermopterotic. The parietal is considerably larger. The dermopterotic is longer but much thinner. The supratemporal is sutured to the parietal and reaches the dermopterotic at only one point. As in Joinvillichthys lindstroemi, the long nuchal horn is supported only by the dermosupraoccipital. The orbitosphenoid and the pleurosphenoid are present in the orbit, just below the frontal, but the basisphenoid is not visible. The toothless premaxilla is longer and narrower. The toothless maxilla also is narrower and more elongate. The lower jaw is composed with the same bones but is longer. The dentary bears two small incisiform teeth and its ventral margin is denticulated. The articulation with the quadrate is located at the level of the posterior border of the orbit. A fragment of a large first infraorbital is preserved on the suture between the prefrontal and the premaxilla. The sclerotic ring is visible in the orbit. The hyomandibula and the preopercle are sutured together. The exposed part of the hyomandibuladermohyomandibula is larger than in Joinvillichthys lindstroemi but still much smaller than the considerably enlarged preopercle. The opercle is broader and comma-shaped, with the sharp end dorsally located. A part of the anterior ceratohyal and two small branchiostegal rays are visible behind the lower jaw. 2. The girdles (Figs 14, 16) The bones of the gigantic pectoral girdle have the same size and shape as in Joinvillichthys lindstroemi. However, two important differences exist. The anterior ventral branch of the cleithrum is lost. No postcleithrum is visible, but that is perhaps due to the fossilization. The pectoral spine is shorter, much broader and is not articulated with the rear of the cleithrum but more anteriorly on its lower margin. A small pelvic girdle is present. Indeed, a part of a vertically oriented pelvic bone is visible under a broken part of the cleithrum. 3. The axial skeleton (Fig. 13) The trunk is fusiform but proportionally deeper than in Joinvillichthys lindstroemi. The axial skeleton is incomplete. Three vertebral segments are missing near the caudal peduncle. There are 16 neural spines (the three missing ones included) before the epichordal series. Only 8 haemal spines are preserved. The total number of haemal spines must be around 12 or 13. The neural and haemal spines are short but broad. The neural and haemal arches surround almost completely the notochord. No ribs are visible. The postcoelomic bone is well developed and backwardly oriented. 4. The dorsal and anal fins (Fig. 13) The short dorsal and anal fins are located at the mid-length of the body. There are 9 pterygiophores and 9 rays in the dorsal fin. The anal fin contains 7 rays, but the number of anal pterygiophores is not determinable. The first dorsal and anal ray is spiny. The other rays are segmented. 5. The caudal skeleton (Fig. 15) The caudal skeleton of the holotype is partly preserved. There are 6 short and broad epichordals and 7 hypochordals. However, one or two anterior hypochordals are missing, so the complete series must be composed of 8 or 9 elements. The fifth preserved hypochordal is strongly enlarged. No urodermal is visible, but the region where theses bones are usually present is not preserved. The caudal fin is double emarginated (Poyato-Ariza & Wenz 2002: fig. 36E) and contains 19 principal segmented caudal rays, the 2 external being pointed and the 17 others branched. There are 6 ventral and at least 4 dorsal procurrent rays. 6. Squamation (Fig. 16) The squamation is the same as in Joinvillichthys lindstroemi. There are 7 spiny scutes in the dorsal ridge and at least 3 spiny scutes in the ventral keel. The two posterior ventral scutes are associated with the ventral margin of the postcoelomic bone. The body scales are slightly ornamented with tubercles. Anteriorly, they are small and flake-like. Posteriorly, there are much larger, irregular and scute-like shaped., Published as part of Taverne, Louis & Capasso, Luigi, 2014, On the " Coccodus " lindstroemi species complex (Pycnodontiformes, Gladiopycnodontidae) from the marine Late Cretaceous of Lebanon, with the description of two new genera, pp. 1-27 in European Journal of Taxonomy 101 on pages 13-19, DOI: 10.5852/ejt.2014.101, http://zenodo.org/record/3838770, {"references":["Hay O. P. 1903. On a collection of Upper Cretaceous fishes from Mount Lebanon, Syria, with descriptions of four new genera and nineteen new species. Bulletin of the American Museum of Natural History 19 (10): 395 - 452.","Poyato-Ariza F. J. & Wenz S. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139 - 248."]}

Joinvillichthys gen. nov. urn:lsid:zoobank.org:act: 6D820D8E-DF5C-41E9-B351-F46F7A9D4119 Type species: Coccodus lindstroemi Davis, 1890 (here designated). Diagnosis Gladiopycnodontid with an elongate prefrontal forming a short rostrum outpacing the lower jaw level. Anterior extremity of the prefrontal acuminate and spiny. Vomer bearing small rounded molariform teeth irregularly ranged. Orbitosphenoid, pleurosphenoid and basisphenoid present. Supratemporal sutured to the rear of the skull. Premaxilla long, broad, toothless and sutured by its upper margin to the prefrontal. Dentary bearing 2 incisiform teeth. Hypertrophied trapezoid preopercle covering the cheek. Large first infraorbital (only known in Joinvillichthys kriweti gen. et sp. nov.). Exposed part of the hyomandibula-dermohyomandibula much smaller than the preopercle. Long nuchal horn with a spiny posterior margin and articulated only on the dermosupraoccipital. Pectoral girdle closely associated to the skull, forming a cephalo-thorax. Cleithrum hypertrophied, with a gigantic posterior ventral process. Hypercleithrum hypertrophied. Well developed posttemporal, with an acuminate posterior extremity. 16 to 17 neural spines, all fused to the neural arches, before the epichordal series. 10 haemal spines before the hypochordal series (only known in Joinvillichthys lindstroemi). Short dorsal fin with 8 or 9 rays. Short anal fin with 7 to 9 rays. 7 to 9 spiny scutes in the dorsal ridge. 3 or 4 scutes in the ventral keel, the last scute or the two last ones associated to the postcoelomic bone. Body completely covered by small, flake-like scales in the abdominal region and by large, scute-like scales in the caudal region. Etymology The generic name is chosen in memory of Lord Jean de Joinville (1224–1317), seneschal of Champagne, who related in his biography of Louis IX the presentation of some Lebanese fossil fishes to this French king at Saïda during the seventh crusade (Gayet et al. 2012: 8). The Greek word ichthys, fish, is added.

Genus Rhinopycnodus gen. nov. urn:lsid:zoobank.org:act: 61F4FDD2-68EC-49EE-B7AB-F36120047FDB Type-species Rhinopycnodus gabriellae gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Greek ris, rinos, the nose. Indeed, the upper jaw of the fish looks like a hog snout when seen in profile. The generic name Pycnodus is added.

Rhinopycnodus gabriellae gen. et sp. nov. Figs 1-8 urn:lsid:zoobank.org:act: D5399E6C-45DF-4D7D-BA77-545D6014BE22 Diagnosis Primitive pycnodontid characterized by a long and broad premaxilla bearing one short and very broad tooth. Elongated head with a long preorbital region. Dentary bearing 2 small incisiform teeth. No temporal fossa. Exoccipital-basioccipital region well visible behind the dermopterotic. Mouth gape obliquely oriented. Preopercle larger than the exposed region of the hyomandibula-dermohyomandibula. Maximum body depth: 67.6 % of the SL. Pectoral fin with 9 rays. Ventral fin with 3 rays. Dorsal fin with 49 pterygiophores. Origin of the dorsal fin behind the dorsal apex and at 76.2 % of the SL. Anal fin with 47 rays and 45 pterygiophores. Origin of the anal fin on the ventral apex and at 63.8 % of the SL. Neural and haemal arches almost completely surrounding the notochord. 32 vertebral elements (neural spines) before the epichordal series. Neural spines 1-8 autogenous. 11 pairs of ribs. Postcoelomic bone reaching the axial skeleton and the ventral margin. 15 haemal spines before the hypochordal series. 7 epichordal and 11 hypochordal elements in the caudal skeleton. Caudal fin with 30 principal rays. Body scales only in the abdominal region. Complete scales ventrally and scale bars dorsally. 19 dorsal ridge scales. First dorsal ridge scale small, triangular in shape and not sutured to the dermosupraoccipital. Some spiny scales in the dorsal ridge and in the ventral keel. 3 pelvic scales. 2 postcloacal scales. Etymology The species name of the new Lebanese fossil fish is dedicated to Mrs. Gabriella di Tota, the co-author’s wife. Formation and locality Marine Upper Cenomanian, Haqel, Lebanon. Holotype and unique specimen Sample CLC S-725, a complete specimen seen by its right side (Figs 1, 2) from Haqel, Lebanon. Total length: 223 mm. Holotype morphometric data The morphometric data are given in % of the standard length (183 mm) of the holotype. Length of the head (opercle included) ……………………………………………………………45.2 % Depth of the head (in the occipital region) ………………………………………………………36.7 % Maximum depth of the body (at the anal fin origin level) ………………………………………67.6 % Prepelvic length …………………………………………………………………………………52.4 % Predorsal length …………………………………………………………………………………76.2 % Basal length of the dorsal fin ……………………………………………………………………39.0 % Preanal length ……………………………………………………………………………………63.8 % Basal length of the anal fin ………………………………………………………………………40.0 % Osteology 1. The skull (Figs 3, 4) The head is high, with the preorbital region much longer than the orbital-postorbital region. The dermal bones of the skull are slightly ornamented with small granulations. The mouth gape is inclined ventrally. The mesethmoid is broad, very long and its upper margin is covered by a pair of long and very narrow prefrontals. The vomer is seen in profile and only six rounded molariform teeth ranged in one rank are visible. The frontal is rather short and not very broad. The posterior margin of the bone is a little enlarged and meets the dermosupraocciptal, the parietal and the small autosphenotic, but not the dermopterotic. Posteriorly, the dermosupraoccipital ends in a short pointed tip. The parietal bears a large posterior brush-like process. There is no temporal fenestra. The dermopterotic is not deepened and is located at the level of the upper border of the orbit. The opisthotic, intercalar, basioccipital and exoccipital are visible behind the dermopterotic and the hyomandibula. A small orbitosphenoid is pressed against the posterior border of the mesethmoid. The parasphenoid is very long and its trabecular region is obliquely oriented. Posteriorly, the parasphenoid reaches the level of the basioccipital. The sensory canals on the braincase are not visible. The quadratic arch contains a well developed quadrate, a large metapterygoid, a large entopterygoid and a small ectopterygoid. The quadrate and the symplectic are both articulated on the lower jaw. The premaxilla is long and very broad. It bears only one short but very broad tooth. The maxilla is small, deeper than long, reniform, toothless and pressed against the premaxilla by its upper margin. When seen in profile, the upper jaw is hog snout-like because of the broadening of the premaxilla and of its tooth. The lower jaw is small and triangular in shape. The dentary is reduced to its ventral branch and it bears two small incisiform teeth. The angular covers a great part of the external face of the mandible. The small articular and the dentary are articulated together. The coronoid process of the prearticular is well marked. Three deep molariform teeth fallen from the prearticular are visible just behind the jaw. The last infraorbital is long, well ossified and forms the posterior and ventral margins of the orbit. No other infraorbital is preserved. The dermosphenotic is lost. The preopercle is the largest bone of the skull, all together deep and broad. The hyomandibula and dermohyomandibula are fused and their exposed part is important but, however, much smaller than the preopercle. The opercle is well developed, with a pointed ventral tip and a broader upper part. No branchiostegal ray is preserved. The anterior ceratohyal is the only preserved part of the hyoid bar. A few hook-like branchial teeth and some branchial filaments are visible under some broken regions of the opercle and preopercle. 2. The girdles (Figs 3-5) Only the ventral part of the posttemporal is preserved. The hypercleithrum (= supracleithrum) is deep, rather broad and not splint-like as usual in Pycnodontiformes. The cleithrum is a large bone with a broad palaform ventral branch and a sinus in its posterior margin to house the pectoral fin, which is short and contains a least 9 rays. The two pelvic bones are vertically oriented. The ventral fins are rather long. Each of them contains 3 rays. The origin of the ventral fins is located a little before the midpoint of the ventral contour. 3. The axial skeleton (Fig. 2) Starting from the caudal region, the vertebral axis progressively elevates to reach the level of the orbit anteriorly. The vertebrae are constituted by only the dorsal and ventral arcocentra. No chordacentrum or autocentrum is visible. The neural and haemal arches surround the notochord almost completely. There are 32 neural spines before the epichordal series and thus 32 vertebral segments before the elements involved in the caudal fin support. The first 17 vertebral segments are abdominal and the following 15 caudal. The anteriormost 8 neural spines are autogenous. The first seven lean on the occipital region of the braincase and the eighth spine is located just above the first ossified but small basidorsal. The basiventrals are strongly reduced in the abdominal region but well developed in the caudal region. There are 11 pairs of long alate ribs and 15 haemal spines before the hypochordal series. The two last pairs of ribs are noticeably shorter than the nine preceding ribs. The first three haemal spines are pressed together and against the postcoelomic bone. The first haemal spine is incomplete and does not reach the axial skeleton. The neural and haemal spines bear an anterior sagittal flange, except for the first five neural and the first three haemal spines. A well developed postzygaphophysis links each neural and haemal arch with the following one. The postcoelomic bone is a long and robust bone dorsally reaching the axial skeleton and ventrally the lower margin of the abdomen. 4. The dorsal and anal fins (Fig. 2) The dorsal fin begins a little behind the dorsal apex. The fin is supported by 49 long and strong pterygiophores. The first seventeen of them have lost the corresponding rays. The last thirty two pterygiophores bear short segmented and branched rays. The first two pterygiophores are broader than the following ones. The anal fin is strip-like in shape (type A2 of Poyato-Ariza & Wenz 2002: fig. 34). The origin of the fin is located at the ventral apex. There are 45 strong pterygiophores bearing 47 rays. The first five pterygiophores abut against the postcoelomic bone and are progressively lengthened from the first to the fifth. The first three rays are reduced to short spines. The other rays are segmented and branched. The very short first pterygiophore supports two little spiny rays and the last pterygiophore two soft rays. 5. The caudal skeleton (Figs 6, 7) There is no caudal peduncle because the dorsal and anal fins reach the caudal fin.The caudal endoskeleton contains 1 urodermal, 7 epichordal and 11 hypochordal elements. The first epichordal neural arch bears a long and narrow neural spine but the length of the neural spines progressively decreases from the first to the seventh epichordal element, which has only a very short neural spine. Some elements in the hypochordal series are broadened but there is no real hypertrophy. This broadening is more important on the eighth and the tenth hypochordal elements than on the other parts of the series. The contour of the caudal fin is double emarginated (Poyato-Ariza & Wenz 2002: fig. 36 E) but the median convex part of the fin is greatly enlarged. There are 30 principal rays, 3 dorsal and 4 ventral procurrent rays. 6. Squamation (Figs 2, 4, 5, 8) The dorsal ridge and ventral keel scales are notably differentiated from the flank scales. There are 19 dorsal ridge scales between the dermosupraoccipital and the origin of the dorsal fin but only the first and the seventh to the tenth are well preserved. The first dorsal scale is small, triangular in shape and located just behind the dermosupraoccipital. The five following scales are badly crushed. Only fragments of the last nine are visible in a fissure of the substratum at the dorsal apex level. Each upper margin of the seventh to tenth dorsal scales bears up to six small spines. The total number of ventral ke el scales is unknown. The ventral keel begins with 3 scales located under the cleithrum (Fig. 4). The first one bears a few very small spines. The second one has a large spine and the third one two large spines. A few ventral keel scales bearing very small spines are visible on the ventral contour between the cleithrum and the pelvic girdle, but some elements of this series are lost because of the taphonomic events. There are 3 pelvic scales associated with the pelvic bones and 2 postcloacal scales are located just before the postcoelomic bone. There are flank scales only in the abdominal region of the body, anterior to the origin of the dorsal and anal fins. In the ventralmost area of the situs viscerum, between the cleithrum and the postcoelomic bone, the scales are complete, thick, deep, broad, slightly ornamented with small tubercules and articulated together. There are 11 rows of these large ventral flank scales. The other body scales are reduced to scale bars. In the dorsal area of the abdominal region the scale bars are badly preserved and only fragments are visible between the neural spines. Scale bars also are associated with the first eight dorsal ridge scales. The scales linked to the eleven other dorsal ridge scales are progressively broader and longer. The first scale of the lateral line is visible beneath the brush-like process of the parietal., Published as part of Taverne, Louis & Capasso, Luigi, 2013, Osteology and relationships of Rhinopycnodus gabriellae gen. et sp. nov. (Pycnodontiformes) from the marine Late Cretaceous of Lebanon, pp. 1-14 in European Journal of Taxonomy 67 on pages 3-11, DOI: 10.5852/ejt.2013.67, http://zenodo.org/record/3827652, {"references":["Poyato-Ariza F. J. & Wenz S. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139 - 248."]}

Genus Monocerichthys gen. nov. urn:lsid:zoobank.org:act: 2D642D4F-2B85-4838-90BD-B967E8345947 Type-species Monocerichthys scheuchzei gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Greek monoker��s, unicorn, and ichthys, fish., Published as part of Taverne, Louis & Capasso, Luigi, 2013, Gladiopycnodontidae, a new family of pycnodontiform fishes from the Late Cretaceous of Lebanon, with the description of three genera, pp. 1-30 in European Journal of Taxonomy 57 on page 11, DOI: 10.5852/ejt.2013.57, http://zenodo.org/record/3822966

Genus Gladiopycnodus gen. nov. urn:lsid:zoobank.org:act: 623891D8-C2AE-4814-91C3-CBE48D098B88 Type-species Gladiopycnodus karami gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Latin gladius, sword, referring to the shape of the pectoral and anal spines. The generic name Pycnodus is added., Published as part of Taverne, Louis & Capasso, Luigi, 2013, Gladiopycnodontidae, a new family of pycnodontiform fishes from the Late Cretaceous of Lebanon, with the description of three genera, pp. 1-30 in European Journal of Taxonomy 57 on page 4, DOI: 10.5852/ejt.2013.57, http://zenodo.org/record/3822966

Genus Rostropycnodus gen. nov. urn:lsid:zoobank.org:act: 66711DD0-8E43-4531-B64A-21071FE34C98 Type-species Rostropycnodus gayeti gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Latin rostrum, rostrum. The generic name Pycnodus is added., Published as part of Taverne, Louis & Capasso, Luigi, 2013, Gladiopycnodontidae, a new family of pycnodontiform fishes from the Late Cretaceous of Lebanon, with the description of three genera, pp. 1-30 in European Journal of Taxonomy 57 on page 19, DOI: 10.5852/ejt.2013.57, http://zenodo.org/record/3822966

Gladiopycnodus karami gen. et sp. nov. Figs 1-6 urn:lsid:zoobank.org:act: BE468D57-6344-4EA5-8F40-2A0646799E2C Diagnosis Gladiopycnodontid with an extremely elongated snout, forming a sword-like rostrum horizontally oriented and greatly outpacing the lower jaw level. Anterior tip of the prefrontal ending in an acuminate point. Frontal long and narrow. Dermosphenotic and dermopterotic partly fused. Oral border of premaxilla and maxilla toothless but bearing very small spines. Five infraorbitals, the first one larger than the others. Large lower jaw horizontally oriented. Large prearticular bearing molariform teeth. Preopercle greatly hypertrophied, with an elongated and denticulated lower margin forming the ventral border of the skull behind the lower jaw. Cleithrum vertically oriented. Strong pectoral spine. Reduced pelvic girdle present. Nuchal horn absent. Long and robust postcoelomic bone. Dorsal fin with short isolated rays forming a series of finlets. Anal fin beginning with an extremely long and strong spine that greatly outpaces the tail level. Only a few short soft anal rays. Caudal fin with a convex posterior border. Large paired dorsal scutes between the skull and the tail. Body completely covered with small flakelike scales irregularly imbricated. Large rounded scales on the caudal peduncle. Two pelvic scutes. One postcloacal scute. No scute on the ventral margin of the body in the caudal region. Etymology The species name of the new Lebanese fossil fish is dedicated to Youssef Bey Karam (1823 - 1889), the famous hero of Lebanon who lived during the time this country was ruled by the Ottoman Empire. Holotype Sample CLC S-393, a complete specimen seen by its right side (Figs 1-2). Total length (the anal spine comprised): 117 mm. Total length (only skull and body): 98 mm. Other material Three other specimens of the new species exist in private collections and, unfortunately, are not accessible for scientific study. However, the authors received good photos of these three specimens and a few anatomical details, not well preserved on the holotype, were revealed by their observation. Formation and locality Marine Upper Cenomanian, Haqel, Lebanon. Holotype morphometric data The morphometric data are given in % of the holotype standard length (91 mm). Length of the head (rostrum included).........................................................70.0 % Depth of the head (in the occipital region)...................................................27.5 % Length of the rostrum (part anterior to the lower jaw level)........................32.0 % Maximum depth of the body (at the pectoral girdle level)...........................33.0 % Depth of the body at the anal fin origin........................................................20.5 % Length of the pectoral spine......................................................................... 21.5 % Predorsal length............................................................................................75.5 % Basal length of the dorsal fin........................................................................25.0 % Preanal length...............................................................................................84.5 % Basal length of the anal fin (spine comprised)............................................. 11.0 % Length of the anal spine............................................................................... 43.5 % Depth of the caudal peduncle..........................................................................6.5 % Osteology 1. The skull (Fig. 3) The head, rostrum comprised, is twice as long as the body. The dermal bones of the skull are ornamented with small tubercles and a few thin ridges on the anterior portion of the prefrontal. The snout is greatly elongated, forming a sword-like rostrum that is horizontally oriented in the same axis as the braincase and the axial skeleton. This rostrum consists of the prefrontals, the premaxillae, the mesethmoid, the parasphenoid and probably the vomer. All these bones are very long. The most anterior parts of the mesethmoid and of the parasphenoid are hidden by the prefrontal and the premaxilla that are connected together. The vomer is hidden by the premaxilla. The pointed anterior tip of the rostrum is only formed by the prefrontal. The olfactive foramen on the mesethmoid is located at the level of the suture between the frontal and the prefrontal. The skull roof comprises the dermosupraoccipital and paired frontals, parietals, dermopterotics and dermosphenotics. The frontal is flat, very long but narrow. Posteriorly, the bone is a little broader and contacts the dermosupraoccipital and the parietal. It also overhangs the well developed triangular autosphenotic. The dermosupraoccipital exhibits a pointed posterior corner but no real horn. This posterior spiny process reaches the first dorsal scute. The parietal is a large bone. It is not possible to say if a brush-like posterior process is present or not on the parietal, the region being hidden by the pectoral girdle and the first dorsal scutes. The zone where a dermocranial fenestra is present in some pycnodontiform fishes is broken on the holotype. However, photos of other specimens clearly show that such a fenestra does not exist in this species. The dermopterotic and the dermosphenotic are smaller bones than the parietal. They are fused together dorsally but remain separated ventrally. The parasphenoid is long and toothless.A very small orbitosphenoid is present just behind the mesethmoid. The other endocranial bones of the braincase are hidden by the preopercle and the hyomandibuladermohyomandibula. The upper jaw is a part of the rostrum and, thus, is located before the lower jaw, which does not participate in the rostrum. The premaxilla is a very long and rather broad bone that is located below the prefrontal and is sutured with it along its upper margin. The maxilla lies under the posterior extremity of the premaxilla. It is a much shorter bone shaped as a lance head with the point anteriorly oriented. The oral border of both bones bear very small spines but there are no true teeth. The lower jaw is large and triangular in shape. The dentary is a small but long bone, reduced to its ventral branch and bearing two incisiform prehensile teeth. The upper part of the anterior margin of the dentary is ornamented with a few very small spines. The prearticular is the largest component of the lower jaw. Its inner face is never visible, neither on the holotype nor on the other specimens. However, some large molariform teeth are visible under a broken part of the prearticular on the photo of one specimen. The angular is as deep as long. The articular is longer but narrow. Both the quadrate and the symplectic articulate with the lower jaw. As usual in pycnodontiform fishes, the quadrate does not possess an ossified quadratic process. The symplectic is a very robust curved bone pressed against the quadrate. A small metapterygoid overhangs the quadrate. The entopterygoid is a large and broad bone as long as the lower jaw. No ectopterygoid is visible. There are five infraorbitals and a sclerotic ring. The ventral part of the first infraorbital is missing on the holotype, but an enlarged triangular-shaped first infraorbital is visible on the photo of one of the other specimens. However, this bone covers only a very small part of the cheek. The four other infraorbitals are well developed but narrower than the first one and more or less reduced to their neurodermic component. The greatly hypertrophied preopercle is by far the largest bone of the skull, all together deep and broad, with a very elongated ventral border that forms the ventral margin of the skull. Dorsally, the preopercle reaches the dermosphenotic and the autosphenotic and exhibits a posterior hollow in which the hyomandibula-dermohyomandibula fits. The opercle is well developed but a lot smaller than the preopercle. It is an ovoid bone with its dorsal and ventral extremities acuminate. The branchiostegal rays are not visible. The hyomandibula and dermohyomandibula are intimately fused in a pyriform-shaped bone of the same size as the opercle and thus much smaller than the preopercle.Within this double composed bone, the small hyomandibula is anteriorly located and has a smooth surface, whereas the larger dermohyomandibula is posteriorly located and has a granulated surface. It is possible that a ventral branch of the hyomandibula is hidden by the preopercle. The hyoid bar is not visible. 2. The girdles (Figs 2, 4-5) The pectoral girdle is pressed against the skull. The dermal bones of the pectoral girdle are ornamented with small tubercles as those on the skull. The cleithrum is by far the largest element of the girdle. It is a very deep, broad and vertically oriented bone. There is a shortening in its upper part that creates a sort of hollow in its anterior margin. The opercle fits in this hollow. The posttemporal is a very small and more or less rounded bone wedged between the parietal, the supraoccipital and the first dorsal scute. The hypercleithrum (= supracleithrum) is larger and amphora-like in shape. The pectoral fin is lost and replaced by a big pointed spine, articulated on the cleithrum. The pelvic girdle is hidden on the holotype because the fossilisation has pressed the pectoral and anal spines against each other. However, on one specimen seen in photo, two small, obliquely oriented pelvic bones are clearly visible between these two spines and some fragments of short pelvic rays appear under the pelvic scutes. 3. The axial skeleton The body is fusiform. It is difficult to describe the axial skeleton in a detailed way, as it is generally more or less covered by the scales and it is not possible to count the vertebral segments. The holotype shows a few fragments of neural and haemal spines. On one of the specimens seen on photo, many scales on the body are lost and we can observe that the notochord is nearly completely surrounded by the neural and haemal arches. The neural and haemal spines are well developed but rather short. Each of them bears a broad anterior wing. The spines are not interlocked together by interdigitating sutures. The presence of ribs is uncertain, the situs viscerum being hidden by the gigantic preopercle, the opercle and the pectoral girdle. The post-coelomic bone is long and very robust. It reaches the vertebral axis dorsally and the inferior border of the fish ventrally. Its broad ventral part is backwardly curved. 4. The dorsal and anal fins (Figs 2, 4) The dorsal fin is composed of about ten very short, branched rays. They rise between the paired dorsal scutes and remain isolated from each other, forming a series of dorsal finlets. In one of the photographed specimens, the last dorsal rays near the tail are not isolated but pressed together. The anal fin begins with a very strong and extremely long spine that extends backward greatly beyond the tail. A few short, branched rays follow this spine. The surface of the spine is ornamented with granulations ranged in regular ranks. The upper border of the spine bears two big and a series of small denticles. The spine is articulated on the post-coelomic bone and on the first pterygiophore, which is longer and separated from the other anal pterygiophores. 5. The caudal skeleton (Fig. 2) The caudal skeleton is covered by scales and thus remains unknown. The small caudal fin is not forked. Its posterior border is convex. The holotype has 17 principal rays, of which the most external dorsal and ventral ones are segmented, pointed and a little shorter than the 15 others, that are segmented and branched. There are 5 dorsal and 5 ventral procurrent rays. 6. Squamation (Figs 5-6) Gladiopycnodus karami gen. et sp. nov. possesses four different types of scutes and scales. The dorsal margin of the fish is covered by large and deep paired scutes that are ornamented with tubercles and some small irregular ridges. The two last scutes are less deep than the more anterior pieces of the series. The holotype bears thirteen pairs of these dorsal scutes. One sample seen on photo has about twenty paired dorsal scutes. This difference in the number of dorsal scutes perhaps reflects an individual variation within the species, but could have also a sexual origin. The ventral margin of the caudal region of the fish is devoid of such scutes. The body is entirely covered by small flake-like scales that are irregularly imbricated. Most scales have a smooth posterior border, but some of them bear a few very small spines on their posterior border. A series of large rounded scales cover the caudal peduncle. They are ornamented with tubercles. Two pelvic scutes and one postcloacal scute are visible on the photo of the specimen that shows the pelvic girdle. A part of their margins is spiny.

Gladiopycnodus karami gen. et sp. nov. Figs 1-6 urn:lsid:zoobank.org:act: BE468D57-6344-4EA5-8F40-2A0646799E2C Diagnosis Gladiopycnodontid with an extremely elongated snout, forming a sword-like rostrum horizontally oriented and greatly outpacing the lower jaw level. Anterior tip of the prefrontal ending in an acuminate point. Frontal long and narrow. Dermosphenotic and dermopterotic partly fused. Oral border of premaxilla and maxilla toothless but bearing very small spines. Five infraorbitals, the first one larger than the others. Large lower jaw horizontally oriented. Large prearticular bearing molariform teeth. Preopercle greatly hypertrophied, with an elongated and denticulated lower margin forming the ventral border of the skull behind the lower jaw. Cleithrum vertically oriented. Strong pectoral spine. Reduced pelvic girdle present. Nuchal horn absent. Long and robust postcoelomic bone. Dorsal fin with short isolated rays forming a series of finlets. Anal fin beginning with an extremely long and strong spine that greatly outpaces the tail level. Only a few short soft anal rays. Caudal fin with a convex posterior border. Large paired dorsal scutes between the skull and the tail. Body completely covered with small flakelike scales irregularly imbricated. Large rounded scales on the caudal peduncle. Two pelvic scutes. One postcloacal scute. No scute on the ventral margin of the body in the caudal region. Etymology The species name of the new Lebanese fossil fish is dedicated to Youssef Bey Karam (1823 - 1889), the famous hero of Lebanon who lived during the time this country was ruled by the Ottoman Empire. Holotype Sample CLC S-393, a complete specimen seen by its right side (Figs 1-2). Total length (the anal spine comprised): 117 mm. Total length (only skull and body): 98 mm. Other material Three other specimens of the new species exist in private collections and, unfortunately, are not accessible for scientific study. However, the authors received good photos of these three specimens and a few anatomical details, not well preserved on the holotype, were revealed by their observation. Formation and locality Marine Upper Cenomanian, Haqel, Lebanon. Holotype morphometric data The morphometric data are given in % of the holotype standard length (91 mm). Length of the head (rostrum included).........................................................70.0 % Depth of the head (in the occipital region)...................................................27.5 % Length of the rostrum (part anterior to the lower jaw level)........................32.0 % Maximum depth of the body (at the pectoral girdle level)...........................33.0 % Depth of the body at the anal fin origin........................................................20.5 % Length of the pectoral spine......................................................................... 21.5 % Predorsal length............................................................................................75.5 % Basal length of the dorsal fin........................................................................25.0 % Preanal length...............................................................................................84.5 % Basal length of the anal fin (spine comprised)............................................. 11.0 % Length of the anal spine............................................................................... 43.5 % Depth of the caudal peduncle..........................................................................6.5 % Osteology 1. The skull (Fig. 3) The head, rostrum comprised, is twice as long as the body. The dermal bones of the skull are ornamented with small tubercles and a few thin ridges on the anterior portion of the prefrontal. The snout is greatly elongated, forming a sword-like rostrum that is horizontally oriented in the same axis as the braincase and the axial skeleton. This rostrum consists of the prefrontals, the premaxillae, the mesethmoid, the parasphenoid and probably the vomer. All these bones are very long. The most anterior parts of the mesethmoid and of the parasphenoid are hidden by the prefrontal and the premaxilla that are connected together. The vomer is hidden by the premaxilla. The pointed anterior tip of the rostrum is only formed by the prefrontal. The olfactive foramen on the mesethmoid is located at the level of the suture between the frontal and the prefrontal. The skull roof comprises the dermosupraoccipital and paired frontals, parietals, dermopterotics and dermosphenotics. The frontal is flat, very long but narrow. Posteriorly, the bone is a little broader and contacts the dermosupraoccipital and the parietal. It also overhangs the well developed triangular autosphenotic. The dermosupraoccipital exhibits a pointed posterior corner but no real horn. This posterior spiny process reaches the first dorsal scute. The parietal is a large bone. It is not possible to say if a brush-like posterior process is present or not on the parietal, the region being hidden by the pectoral girdle and the first dorsal scutes. The zone where a dermocranial fenestra is present in some pycnodontiform fishes is broken on the holotype. However, photos of other specimens clearly show that such a fenestra does not exist in this species. The dermopterotic and the dermosphenotic are smaller bones than the parietal. They are fused together dorsally but remain separated ventrally. The parasphenoid is long and toothless.A very small orbitosphenoid is present just behind the mesethmoid. The other endocranial bones of the braincase are hidden by the preopercle and the hyomandibuladermohyomandibula. The upper jaw is a part of the rostrum and, thus, is located before the lower jaw, which does not participate in the rostrum. The premaxilla is a very long and rather broad bone that is located below the prefrontal and is sutured with it along its upper margin. The maxilla lies under the posterior extremity of the premaxilla. It is a much shorter bone shaped as a lance head with the point anteriorly oriented. The oral border of both bones bear very small spines but there are no true teeth. The lower jaw is large and triangular in shape. The dentary is a small but long bone, reduced to its ventral branch and bearing two incisiform prehensile teeth. The upper part of the anterior margin of the dentary is ornamented with a few very small spines. The prearticular is the largest component of the lower jaw. Its inner face is never visible, neither on the holotype nor on the other specimens. However, some large molariform teeth are visible under a broken part of the prearticular on the photo of one specimen. The angular is as deep as long. The articular is longer but narrow. Both the quadrate and the symplectic articulate with the lower jaw. As usual in pycnodontiform fishes, the quadrate does not possess an ossified quadratic process. The symplectic is a very robust curved bone pressed against the quadrate. A small metapterygoid overhangs the quadrate. The entopterygoid is a large and broad bone as long as the lower jaw. No ectopterygoid is visible. There are five infraorbitals and a sclerotic ring. The ventral part of the first infraorbital is missing on the holotype, but an enlarged triangular-shaped first infraorbital is visible on the photo of one of the other specimens. However, this bone covers only a very small part of the cheek. The four other infraorbitals are well developed but narrower than the first one and more or less reduced to their neurodermic component. The greatly hypertrophied preopercle is by far the largest bone of the skull, all together deep and broad, with a very elongated ventral border that forms the ventral margin of the skull. Dorsally, the preopercle reaches the dermosphenotic and the autosphenotic and exhibits a posterior hollow in which the hyomandibula-dermohyomandibula fits. The opercle is well developed but a lot smaller than the preopercle. It is an ovoid bone with its dorsal and ventral extremities acuminate. The branchiostegal rays are not visible. The hyomandibula and dermohyomandibula are intimately fused in a pyriform-shaped bone of the same size as the opercle and thus much smaller than the preopercle.Within this double composed bone, the small hyomandibula is anteriorly located and has a smooth surface, whereas the larger dermohyomandibula is posteriorly located and has a granulated surface. It is possible that a ventral branch of the hyomandibula is hidden by the preopercle. The hyoid bar is not visible. 2. The girdles (Figs 2, 4-5) The pectoral girdle is pressed against the skull. The dermal bones of the pectoral girdle are ornamented with small tubercles as those on the skull. The cleithrum is by far the largest element of the girdle. It is a very deep, broad and vertically oriented bone. There is a shortening in its upper part that creates a sort of hollow in its anterior margin. The opercle fits in this hollow. The posttemporal is a very small and more or less rounded bone wedged between the parietal, the supraoccipital and the first dorsal scute. The hypercleithrum (= supracleithrum) is larger and amphora-like in shape. The pectoral fin is lost and replaced by a big pointed spine, articulated on the cleithrum. The pelvic girdle is hidden on the holotype because the fossilisation has pressed the pectoral and anal spines against each other. However, on one specimen seen in photo, two small, obliquely oriented pelvic bones are clearly visible between these two spines and some fragments of short pelvic rays appear under the pelvic scutes. 3. The axial skeleton The body is fusiform. It is difficult to describe the axial skeleton in a detailed way, as it is generally more or less covered by the scales and it is not possible to count the vertebral segments. The holotype shows a few fragments of neural and haemal spines. On one of the specimens seen on photo, many scales on the body are lost and we can observe that the notochord is nearly completely surrounded by the neural and haemal arches. The neural and haemal spines are well developed but rather short. Each of them bears a broad anterior wing. The spines are not interlocked together by interdigitating sutures. The presence of ribs is uncertain, the situs viscerum being hidden by the gigantic preopercle, the opercle and the pectoral girdle. The post-coelomic bone is long and very robust. It reaches the vertebral axis dorsally and the inferior border of the fish ventrally. Its broad ventral part is backwardly curved. 4. The dorsal and anal fins (Figs 2, 4) The dorsal fin is composed of about ten very short, branched rays. They rise between the paired dorsal scutes and remain isolated from each other, forming a series of dorsal finlets. In one of the photographed specimens, the last dorsal rays near the tail are not isolated but pressed together. The anal fin begins with a very strong and extremely long spine that extends backward greatly beyond the tail. A few short, branched rays follow this spine. The surface of the spine is ornamented with granulations ranged in regular ranks. The upper border of the spine bears two big and a series of small denticles. The spine is articulated on the post-coelomic bone and on the first pterygiophore, which is longer and separated from the other anal pterygiophores. 5. The caudal skeleton (Fig. 2) The caudal skeleton is covered by scales and thus remains unknown. The small caudal fin is not forked. Its posterior border is convex. The holotype has 17 principal rays, of which the most external dorsal and ventral ones are segmented, pointed and a little shorter than the 15 others, that are segmented and branched. There are 5 dorsal and 5 ventral procurrent rays. 6. Squamation (Figs 5-6) Gladiopycnodus karami gen. et sp. nov. possesses four different types of scutes and scales. The dorsal margin of the fish is covered by large and deep paired scutes that are ornamented with tubercles and some small irregular ridges. The two last scutes are less deep than the more anterior pieces of the series. The holotype bears thirteen pairs of these dorsal scutes. One sample seen on photo has about twenty paired dorsal scutes. This difference in the number of dorsal scutes perhaps reflects an individual variation within the species, but could have also a sexual origin. The ventral margin of the caudal region of the fish is devoid of such scutes. The body is entirely covered by small flake-like scales that are irregularly imbricated. Most scales have a smooth posterior border, but some of them bear a few very small spines on their posterior border. A series of large rounded scales cover the caudal peduncle. They are ornamented with tubercles. Two pelvic scutes and one postcloacal scute are visible on the photo of the specimen that shows the pelvic girdle. A part of their margins is spiny., Published as part of Taverne, Louis & Capasso, Luigi, 2013, Gladiopycnodontidae, a new family of pycnodontiform fishes from the Late Cretaceous of Lebanon, with the description of three genera, pp. 1-30 in European Journal of Taxonomy 57 on pages 5-10, DOI: 10.5852/ejt.2013.57, http://zenodo.org/record/3822966

Rostropycnodus gayeti gen. et sp. nov. Figs 15-21 urn:lsid:zoobank.org:act: 89707CF2-AF8E-450E-8D6F-16A0B02EDE93 Diagnosis Gladiopycnodontid with a very long snout, forming a rostrum anteriorly outpacing the lower jaw level (Fig. 18). Anterior tip of the rostrum formed by both the prefrontal and the premaxilla.Anterior extremity of the prefrontal bearing four small spines. Frontal bearing one long pointed horn. Dermosupraoccipital forming a nuchal process. Small lower jaw. Preopercle much larger than the exposed part of the hyomandibula-dermohyomandibula. Posttemporal articulated with the nuchal process. Hypertrophied cleithrum with a long and broad ventral branch. Short and broad pectoral spine. Two large postcleithra. Nuchal horn absent. 17 neural spines and 17 haemal spines before the epichordal and hypochordal series. Dorsal fin with 6 pterygiophores. Anal fin with a long and strong initial spine and 4 short soft rays. Caudal skeleton with 6 epichordal and 7 hypochordal elements. Last three hypochordal elements moderately broadened. Caudal fin with 21 rays and a convex posterior margin. Body entirely covered by large, tuberculated scales with a spiny anterior border. Etymology The species name of the new gladiopycnodontid is dedicated to Mireille Gayet (Lyon), the well known French palaeontologist, who has greatly improved our knowledge of the Late Cretaceous ichthyofauna of Lebanon. Material examined Holotype Sample CLC S-608a, b, the two faces of a nearly complete specimen (Fig. 15). The tip of the rostrum and the caudal fin are missing. Total length: 47.3 mm. Paratypes Sample CLC S-337, a complete specimen (Fig.16). The body and the tail are severely crushed. Total length: 59.8 mm. Sample CLC S-595, a nearly complete specimen (Fig.17). The tip of the rostrum is missing. Total length: 58.6 mm. Formation and locality Marine Upper Cenomanian, Haqel, Lebanon. Holotype morphometric data Due to the incompleteness of the available specimens, no accurate morphological data can be provided and the reconstruction is therefore composite. Osteology 1. The skull (Fig. 19) The head, rostrum and opercular series included, is twice as long as the body. The dermal bones of the skull are ornamented with a few thin ridges and many tubercles. The snout is elongated, forming a long rostrum that is, however, shorter than in Gladiopycnodus karami gen. et sp. nov. The preorbital length, rostrum included, represents 59.5 % of the total length of the skull. The rostrum consists of the prefrontals, the premaxillae, the mesethmoid and the parasphenoid. Only the most posterior parts of the mesethmoid and of the parasphenoid are visible. Both the prefrontal and the premaxilla form the tip of the rostrum. The long and broad prefrontal bears four small spines on its anterior tip as seen on sample CLC S-337. The upper margin of the prefrontal is ornamented with small spines. The vomer remains hidden by the premaxilla. A vomer with very small molariform teeth is present in the family, as shown in some other gladiopycnodontid genera. The frontal and the dermosupraoccipital are the main bones of the skull roof. The frontal is short but broad. Anteriorly, it does not go beyond the orbit. The frontal bears a long pointed horn with a large basis and of which the anterior and posterior borders are spiny. The dermosupraoccipital is enlarged and forms a long and broad nuchal process. The autosphenotic, parietal and dermopterotic are rather small bones. The parietal does not bear a brush-like process. The large supratemporal is pressed against the dermosupraoccipital. The parasphenoid is long and toothless. A well developed orbitosphenoid is articulated on the rear of the mesethmoid. The other endocranial bones of the braincase are hidden by the hyomandibuladermohyomandibula, the opercle and the hypercleithrum. The toothless upper jaw is a part of the rostrum. The premaxilla is very long and rather narrow but, however, a little broader anteriorly than posteriorly. It is located under the prefrontal and is sutured with it all along its upper margin. The maxilla is small, considerably shorter than the premaxilla, lance headshaped and posteriorly located. The lower jaw is small, triangular in shape and, by far, does not reach the anterior extremity of the rostrum. The short dentary bone, well visible on specimen CLC S-608, is reduced to its ventral branch. The teeth are not preserved. The other bones of the mandible are too crushed to allow adequate description. The bones of the palatoquadrate arch are hidden by the preopercle. A large dermosphenotic is the only part preserved of the infraorbital series. The exposed part of the hyomandibula-dermohyomandibula is much smaller than the large preopercle that is, however, not as hypertrophied as in Gladiopycnodus karami gen. et sp. nov. The preopercle is separated from the lower margin of the fish by the cleithrum, another difference from G. karami gen. et sp. nov.. The small opercle is a deep ovoid bone with an acuminate ventral extremity. The hyoid bar and the branchiostegal rays are not visible. A fragment of a branchial bone and a few short nail-shaped branchiospines are preserved in the orbit of specimen S-337. 2. The girdles (Figs 18-19) The massive pectoral girdle is intimately associated with the skull, forming a sort of cephalo-thorax. The posttemporal is well developed and located just behind the hypercleithrum. The hypercleithrum (= supracleithrum) is a large triangular bone with some spines on its posterior border. The cleithrum is really gigantic, with a very broad dorsal branch and a very long, broad and pointed ventral branch reaching the lower jaw level. There are two large postcleithra well visible on sample CLC S-337 and located behind the cleithrum. The pectoral fin is replaced by a short but very broad spine that is partially fused with the cleithrum. Two small pelvic bones are preserved between the cleithrum and the post-coelomic bone on specimen CLC S-595, but the fin rays are lost. 3. The axial skeleton (Fig. 18) The body is fusiform. Holotype CLC S-608a, b shows a complete vertebral column. The notochord is incompletely surrounded by the neural and haemal arches, which are reduced. The neural and haemal spines are well developed. There are 17 neural spines before the epichordal series and 17 haemal spines before the hypochordal series. The first ten neural spines are thin. The seven other spines are shorter but also broader. The first three haemal spines are rather short. Their length progressively increases from the fourth to sixth one and then decreases from the seventh to the seventeenth. The neural and haemal spines are not interlocked together by interdigitating sutures. The presence of ribs is uncertain, the situs viscerum being hidden by the gigantic cleithrum. The post-coelomic bone is moderately long, robust, extremely broad and backwardly curved. Dorsally it reaches the vertebral axis and ventrally the inferior border of the fish. 4. The dorsal and anal fins (Fig. 20) The dorsal fin is short. Holotype CLC S-608b shows 6 dorsal pterygiophores, largely separated from each other. The dorsal rays are lost but, considering the spacing between the pterygiophores, they surely formed a series of isolated finlets as in Gladiopycnodus karami gen. et sp. nov. The anal fin begins with a large and strong spine. This spine has spiny dorsal and ventral margins, an acuminate posterior tip and a very broad basis articulated on the post-coelomic bone and the first anal pterygiophore, which is much longer than the following ones. The pointed tip of the spine reaches the level of the tail but does not outpace the caudal fin as in G. karami gen. et sp. nov. Four other small anal pterygiophores, bearing four short rays, are visible on holotype CLC S-608a. 5. The caudal skeleton (Fig. 21A) The caudal skeleton is rather well preserved on holotype CLC S-608a, b. There are 6 epichordal elements and 7 hypochordal elements. The last three hypochordal pieces are moderately broadened. No urodermal is visible. Paratype CLC S-595 possesses a more or less complete caudal fin, but essentially preserved as a print. There are 21 rays, but it is not possible to distinguish the procurrent from the principal rays. The fin has a convex distal border (Poyato-Ariza & Wenz 2002: fig. 36 B). 6. Squamation (Fig. 21B) The body is entirely covered by large deep scales that are imbricated. Their entire surface is ornamented with well developed tubercles and their anterior border is often irregularly spiny, as clearly seen on paratype CLC S-595. The first scales of the dorsal margin of the fish are of the same type, but still larger than those on the flanks. Generally, on the other samples, only the tubercles are preserved, but not the outlines of the scales. Two large, irregularly shaped pelvic scales are visible below the pelvic bones on paratype CLC S-595., Published as part of Taverne, Louis & Capasso, Luigi, 2013, Gladiopycnodontidae, a new family of pycnodontiform fishes from the Late Cretaceous of Lebanon, with the description of three genera, pp. 1-30 in European Journal of Taxonomy 57 on pages 19-26, DOI: 10.5852/ejt.2013.57, http://zenodo.org/record/3822966, {"references":["Poyato-Ariza F. J. & Wenz S. 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1): 139 - 248."]}

Genus Gladiopycnodus gen. nov. urn:lsid:zoobank.org:act: 623891D8-C2AE-4814-91C3-CBE48D098B88 Type-species Gladiopycnodus karami gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Latin gladius, sword, referring to the shape of the pectoral and anal spines. The generic name Pycnodus is added.

Genus Rostropycnodus gen. nov. urn:lsid:zoobank.org:act: 66711DD0-8E43-4531-B64A-21071FE34C98 Type-species Rostropycnodus gayeti gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Latin rostrum, rostrum. The generic name Pycnodus is added.

Rostropycnodus gayeti gen. et sp. nov. Figs 15-21 urn:lsid:zoobank.org:act: 89707CF2-AF8E-450E-8D6F-16A0B02EDE93 Diagnosis Gladiopycnodontid with a very long snout, forming a rostrum anteriorly outpacing the lower jaw level (Fig. 18). Anterior tip of the rostrum formed by both the prefrontal and the premaxilla.Anterior extremity of the prefrontal bearing four small spines. Frontal bearing one long pointed horn. Dermosupraoccipital forming a nuchal process. Small lower jaw. Preopercle much larger than the exposed part of the hyomandibula-dermohyomandibula. Posttemporal articulated with the nuchal process. Hypertrophied cleithrum with a long and broad ventral branch. Short and broad pectoral spine. Two large postcleithra. Nuchal horn absent. 17 neural spines and 17 haemal spines before the epichordal and hypochordal series. Dorsal fin with 6 pterygiophores. Anal fin with a long and strong initial spine and 4 short soft rays. Caudal skeleton with 6 epichordal and 7 hypochordal elements. Last three hypochordal elements moderately broadened. Caudal fin with 21 rays and a convex posterior margin. Body entirely covered by large, tuberculated scales with a spiny anterior border. Etymology The species name of the new gladiopycnodontid is dedicated to Mireille Gayet (Lyon), the well known French palaeontologist, who has greatly improved our knowledge of the Late Cretaceous ichthyofauna of Lebanon. Material examined Holotype Sample CLC S-608a, b, the two faces of a nearly complete specimen (Fig. 15). The tip of the rostrum and the caudal fin are missing. Total length: 47.3 mm. Paratypes Sample CLC S-337, a complete specimen (Fig.16). The body and the tail are severely crushed. Total length: 59.8 mm. Sample CLC S-595, a nearly complete specimen (Fig.17). The tip of the rostrum is missing. Total length: 58.6 mm. Formation and locality Marine Upper Cenomanian, Haqel, Lebanon. Holotype morphometric data Due to the incompleteness of the available specimens, no accurate morphological data can be provided and the reconstruction is therefore composite. Osteology 1. The skull (Fig. 19) The head, rostrum and opercular series included, is twice as long as the body. The dermal bones of the skull are ornamented with a few thin ridges and many tubercles. The snout is elongated, forming a long rostrum that is, however, shorter than in Gladiopycnodus karami gen. et sp. nov. The preorbital length, rostrum included, represents 59.5 % of the total length of the skull. The rostrum consists of the prefrontals, the premaxillae, the mesethmoid and the parasphenoid. Only the most posterior parts of the mesethmoid and of the parasphenoid are visible. Both the prefrontal and the premaxilla form the tip of the rostrum. The long and broad prefrontal bears four small spines on its anterior tip as seen on sample CLC S-337. The upper margin of the prefrontal is ornamented with small spines. The vomer remains hidden by the premaxilla. A vomer with very small molariform teeth is present in the family, as shown in some other gladiopycnodontid genera. The frontal and the dermosupraoccipital are the main bones of the skull roof. The frontal is short but broad. Anteriorly, it does not go beyond the orbit. The frontal bears a long pointed horn with a large basis and of which the anterior and posterior borders are spiny. The dermosupraoccipital is enlarged and forms a long and broad nuchal process. The autosphenotic, parietal and dermopterotic are rather small bones. The parietal does not bear a brush-like process. The large supratemporal is pressed against the dermosupraoccipital. The parasphenoid is long and toothless. A well developed orbitosphenoid is articulated on the rear of the mesethmoid. The other endocranial bones of the braincase are hidden by the hyomandibuladermohyomandibula, the opercle and the hypercleithrum. The toothless upper jaw is a part of the rostrum. The premaxilla is very long and rather narrow but, however, a little broader anteriorly than posteriorly. It is located under the prefrontal and is sutured with it all along its upper margin. The maxilla is small, considerably shorter than the premaxilla, lance headshaped and posteriorly located. The lower jaw is small, triangular in shape and, by far, does not reach the anterior extremity of the rostrum. The short dentary bone, well visible on specimen CLC S-608, is reduced to its ventral branch. The teeth are not preserved. The other bones of the mandible are too crushed to allow adequate description. The bones of the palatoquadrate arch are hidden by the preopercle. A large dermosphenotic is the only part preserved of the infraorbital series. The exposed part of the hyomandibula-dermohyomandibula is much smaller than the large preopercle that is, however, not as hypertrophied as in Gladiopycnodus karami gen. et sp. nov. The preopercle is separated from the lower margin of the fish by the cleithrum, another difference from G. karami gen. et sp. nov.. The small opercle is a deep ovoid bone with an acuminate ventral extremity. The hyoid bar and the branchiostegal rays are not visible. A fragment of a branchial bone and a few short nail-shaped branchiospines are preserved in the orbit of specimen S-337. 2. The girdles (Figs 18-19) The massive pectoral girdle is intimately associated with the skull, forming a sort of cephalo-thorax. The posttemporal is well developed and located just behind the hypercleithrum. The hypercleithrum (= supracleithrum) is a large triangular bone with some spines on its posterior border. The cleithrum is really gigantic, with a very broad dorsal branch and a very long, broad and pointed ventral branch reaching the lower jaw level. There are two large postcleithra well visible on sample CLC S-337 and located behind the cleithrum. The pectoral fin is replaced by a short but very broad spine that is partially fused with the cleithrum. Two small pelvic bones are preserved between the cleithrum and the post-coelomic bone on specimen CLC S-595, but the fin rays are lost. 3. The axial skeleton (Fig. 18) The body is fusiform. Holotype CLC S-608a, b shows a complete vertebral column. The notochord is incompletely surrounded by the neural and haemal arches, which are reduced. The neural and haemal spines are well developed. There are 17 neural spines before the epichordal series and 17 haemal spines before the hypochordal series. The first ten neural spines are thin. The seven other spines are shorter but also broader. The first three haemal spines are rather short. Their length progressively increases from the fourth to sixth one and then decreases from the seventh to the seventeenth. The neural and haemal spines are not interlocked together by interdigitating sutures. The presence of ribs is uncertain, the situs viscerum being hidden by the gigantic cleithrum. The post-coelomic bone is moderately long, robust, extremely broad and backwardly curved. Dorsally it reaches the vertebral axis and ventrally the inferior border of the fish. 4. The dorsal and anal fins (Fig. 20) The dorsal fin is short. Holotype CLC S-608b shows 6 dorsal pterygiophores, largely separated from each other. The dorsal rays are lost but, considering the spacing between the pterygiophores, they surely formed a series of isolated finlets as in Gladiopycnodus karami gen. et sp. nov. The anal fin begins with a large and strong spine. This spine has spiny dorsal and ventral margins, an acuminate posterior tip and a very broad basis articulated on the post-coelomic bone and the first anal pterygiophore, which is much longer than the following ones. The pointed tip of the spine reaches the level of the tail but does not outpace the caudal fin as in G. karami gen. et sp. nov. Four other small anal pterygiophores, bearing four short rays, are visible on holotype CLC S-608a. 5. The caudal skeleton (Fig. 21A) The caudal skeleton is rather well preserved on holotype CLC S-608a, b. There are 6 epichordal elements and 7 hypochordal elements. The last three hypochordal pieces are moderately broadened. No urodermal is visible. Paratype CLC S-595 possesses a more or less complete caudal fin, but essentially preserved as a print. There are 21 rays, but it is not possible to distinguish the procurrent from the principal rays. The fin has a convex distal border (Poyato-Ariza & Wenz 2002: fig. 36 B). 6. Squamation (Fig. 21B) The body is entirely covered by large deep scales that are imbricated. Their entire surface is ornamented with well developed tubercles and their anterior border is often irregularly spiny, as clearly seen on paratype CLC S-595. The first scales of the dorsal margin of the fish are of the same type, but still larger than those on the flanks. Generally, on the other samples, only the tubercles are preserved, but not the outlines of the scales. Two large, irregularly shaped pelvic scales are visible below the pelvic bones on paratype CLC S-595.

Genus Monocerichthys gen. nov. urn:lsid:zoobank.org:act: 2D642D4F-2B85-4838-90BD-B967E8345947 Type-species Monocerichthys scheuchzei gen. et sp. nov. (by monotypy) Diagnosis As for the species (monospecific genus). Etymology From the Greek monokerôs, unicorn, and ichthys, fish.