taxonID	type	description	language	source
03C83000B577FFB7FC8A03F3FC88FE6B.taxon	diagnosis	Diagnosis (emended from Prokofiev, 2005): Large-sized paralepidids (most of the specimens reach about 60 cm SL) characterized by a rigid trunk with opposing dorsal and anal-fins posteriorly displaced on the posterior half of the body; dorsal-fin origin located on or slightly behind the vertical of the anal-fin origin; at least 18 dorsal-fin rays; high number of vertebrae (70 – 112); caudal vertebrae 25 – 40 % of the total number; extended series of epineurals and epipleurals reaching the caudal region; body covered by small scales or totally naked; large and dorsoventrally symmetrical caudal fin; (apparent) absence of adipose fin. I n c l u d e d g e n e r a: † H o l o s t e u s A g a s s i z, 1 8 3 5, † Pavlovichthys Menner, 1948. Remarks: The subfamily † Holosteinae was erected by Prokofiev (2005) based on putatively unique synapomorphies shared by † H. esocinus Agassiz, 1835, from the Eocene of Monte Bolca and † ‘ Holosteus’ mariae (Menner, 1948) from the lower Oligocene deposits of Caucasus (Russia) and Carpathians (Romania). According to Prokofiev (2005), these features include a lower jaw joint located in front of the level of the anterior margin of orbit, dorsal-fin origin located behind the anal-fin origin, about 70 – 100 vertebrae, at least ten branchiostegal rays, large fang-like teeth on the dentary and about 25 rays in both anal and pelvic fins. However, the lack of a comprehensive revision of the Eocene species from Monte Bolca prevented the recognition of the actual nature of their relationships. The exquisitely preserved specimens from the Eocene of Monte Bolca described herein are certainly related to the Oligocene taxon and clearly belong to the subfamily † Holosteinae, being characterized by a rigid trunk with both median fins displaced on the posterior half of the body, a dorsal fin with at least 18 rays whose origin is located slightly behind the vertical of that of the anal fin and a high number of vertebrae (70 – 110). Three additional synapomorphies were also recognized, including caudal vertebrae of 25 – 40 % of the total number, intermuscular bones extending posteriorly and reaching the caudal region and (apparent) absence of an adipose fin. A number of features, however, provide robust evidence of the separate generic status of the Eocene and Oligocene species (see below).	en	Marramà, Giuseppe, Carnevale, Giorgio (2017): Morphology, relationships and palaeobiology of the Eocene barracudina † Holosteus esocinus (Aulopiformes: Paralepididae) from Monte Bolca, Italy. Zoological Journal of the Linnean Society 181 (1): 209-228, DOI: 10.1093/zoolinnean/zlw029, URL: http://academic.oup.com/zoolinnean/article/181/1/209/3782581/Morphology-relationships-and-palaeobiology-of-the
03C83000B576FFB7FED70374FAD9FCB8.taxon	materials_examined	Type species: † H. esocinus Agassiz, 1835. Diagnosis: Holosteine paralepidid with very elongated body and fineness ratio of 15.0 – 16.0; head is of about five to six times SL; upper jaw terminating below the ventral margin of the orbit; prominent, non-ossified and horizontally directed projection forming the anterior tip of the lower jaw; articular – quadrate joint located below the posterior margin of the orbit; eight branchiostegal rays; small recurved teeth on premaxilla; large, thin and pointed teeth on palate and lower jaw without corrugate pattern or serrate margins; epineurals and epipleurals extending throughout the vertebral column, being thicker and more ossified in the caudal region; most epineurals bifid or trifid proximally; 106 – 112 vertebrae of which about 30 % are caudal; dorsal fin with 18 – 20 rays; about 19 – 22 anal-fin rays; about 10 – 12 pelvic-fin rays; caudal fin with 19 – 20 principal rays and about 10 – 12 dorsal and ventral procurrent rays; five or six autogenous hypurals, of which the first two appear to be partially fused; proximal ends of the epaxial caudal-fin rays that extensively overlap the associated hypurals; bony fulcral scales just in front of the upper and ventral caudal-fin lobes; body completely naked, with a broad, uniform, mid-dorsal brown-pigmented band along the back. Remarks: † Holosteus esocinus was created by Agassiz (1835) based on a nearly complete specimen in the collection of the MNHN, Paris (F. Bol 175). Recently, Prokofiev (2005) provided a re-examination of the lower Oligocene paralepidids from the Caucasus and Carpathians. Following the taxonomic interpretations of Daniltshenko (1960), Gorbach (1961) and Constantin (2001), Prokofiev (2005) assigned to the genus † Holosteus, the Oligocene barracudina species † Pavlovichthys mariae Menner, 1948, also providing a diagnosis of the genus. This Oligocene species differs from † H. esocinus by having the articular – quadrate joint located in front of the anterior margin of the orbit, large fang-like teeth with corrugate pattern on lower jaw, more than eight branchiostegal rays, first and second hypurals not fused and anal and pelvic fins with about 25 rays. Our observations clearly indicate that the generic diagnosis provided by Prokofiev (2005) for the genus Holosteus must be restricted to the Oligocene taxon, which, as a consequence, should be referred to a separate genus (see above). The Oligocene species † ‘ Holosteus ’ mariae is therefore reassigned to the genus † Pavlovichthys, following the principle of priority of the International Commission on Zoological Nomenclature (ICZN, 1999). In his analysis, Prokofiev (2005) indicated the generic name † Xiphopterus, erected by Agassiz (1833 – 1844) for † Esox falcatus Volta, 1796, from the Eocene of Monte Bolca as a probable synonym of † Holosteus. The examination of the holotype of † Xiphopterus falcatus housed in the MNHN, Paris (F. Bol 544) allows us to exclude such a hypothesis, considering that except for the similar elongated body, none of the diagnostic characters of the Paralepididae can be recognized in this fish.	en	Marramà, Giuseppe, Carnevale, Giorgio (2017): Morphology, relationships and palaeobiology of the Eocene barracudina † Holosteus esocinus (Aulopiformes: Paralepididae) from Monte Bolca, Italy. Zoological Journal of the Linnean Society 181 (1): 209-228, DOI: 10.1093/zoolinnean/zlw029, URL: http://academic.oup.com/zoolinnean/article/181/1/209/3782581/Morphology-relationships-and-palaeobiology-of-the
03C83000B576FFBEFC7202E7FE21FB11.taxon	description	(FIGS 1 – 6)	en	Marramà, Giuseppe, Carnevale, Giorgio (2017): Morphology, relationships and palaeobiology of the Eocene barracudina † Holosteus esocinus (Aulopiformes: Paralepididae) from Monte Bolca, Italy. Zoological Journal of the Linnean Society 181 (1): 209-228, DOI: 10.1093/zoolinnean/zlw029, URL: http://academic.oup.com/zoolinnean/article/181/1/209/3782581/Morphology-relationships-and-palaeobiology-of-the
03C83000B576FFBEFC7202E7FE21FB11.taxon	materials_examined	Holotype: MNHN F. Bol 175, nearly complete articulated skeleton (Fig. 1 A), 347.2 mm SL. Referred material: MCSNV IG. 23601 and IG. 23602, nearly complete articulated skeleton, in part and counterpart (Fig. 1 B, C), 499.4 mm SL; MGPUP 11611 / 2, nearly complete articulated skeleton, in part and counterpart (Fig. 1 D, E), 541.0 mm SL; MCSNV B 5 / T 98, incomplete articulated specimen lacking of part of the abdominal and cranial regions, in part and counterpart (Fig. 2 A, B); MCSNV VI. N 41, partially complete articulated specimen (Fig. 2 C), 644.1 mm SL. Type locality and horizon: Monte Bolca locality, Pesciara site; early Eocene, late Ypresian, middle Cuisian, SBZ 11, Alveolina dainelli Zone (see Papazzoni et al., 2014). Diagnosis: As for the genus. Description: Counts and measurements for † H. esocinus are provided in Tables 1 and 2. The body is laterally compressed and considerably elongated and slender (Figs 1, 2); it is characterized by a remarkably high fineness ratio (15.0 – 16.0), which is lower only than that of the extremely elongated Stemonosudis. The head is elongated and nearly triangular in lateral outline, and its length is contained between five to six times in SL. The snout is pointed and long, measuring c. 50 % of the head length. The mouth is large, terminal, with a lower jaw projecting well anteriorly to the tip of the upper jaw through a non-ossified prominent and horizontally directed pointed projection. Dorsal and anal fins are displaced to the posterior half of the body, opposite to each other; the dorsal-fin origin is located slightly behind the vertical through the anal-fin origin. The adipose fin appears to be absent. The caudal peduncle is narrow and short (about 9 – 12 % SL). The caudal fin is large, symmetrical and forked, with a concave posterior margin. † Holosteus esocinus is totally naked; lateral-line scales seem to be also absent. The description of the skeletal structures of the neurocranium is primarily based on the specimens MCSNV IG. 23601 / 2 and MGPUP 11611 / 2, in which this region is largely exposed and well preserved (Figs 3 A, B, 4 A, B), except for the otic region that is largely incomplete and difficult to interpret. The neurocranium is extremely elongated and nearly triangular in outline. The paired frontals occupy about half of the skull roof length; each frontal articulates anteriorly with the mesethmoid, ventrally with the orbitosphenoid, anteroventrally with the lateral ethmoid, posteriorly with the parietal and posteroventrally with the pterosphenotic and the sphenotic. The parietals are irregular in shape. The supraoccipital forms the posteromedial end of the neurocranium. As in extant paralepidids (Rofen, 1966), the parietals are not fully separated by the supraoccipital (Fig. 4 A, B). The sphenotic is subrectangular in outline, devoid of an anterior process. The orbitosphenoid and pterosphenoid form the dorsal and posterior walls of the orbit, respectively. The epioccipital occupies the dorsolateral part of the posterior surface of the neurocranium. The exoccipitals are badly crushed in all the examined specimens, and their morphology is not clearly recognizable. The parasphenoid is slender and almost Note: Due to the inadequate preservation of the fossil, lacking most of the cranial and abdominal portions of the body, the data for the specimen MCSNV B 5 / T 98 are not reported in the table. Figure 3. † Holosteus esocinus Agassiz, 1835. Reconstructions of the head based on the paratype. (A) MCSNV IG. 23602; (B) MCSNV IG. 23601. Scale bars, 10 mm. Abbreviations: aa, anguloarticular; ac, anterior ceratohyal; br, branchiostegal rays; cl, cleithrum; co, coracoid; de, dentary; ect, ectopterygoid; end, endopterygoid; epo, epioccipital; fe, premaxillary fenestra; fr, frontal; hym, hyomandibula; iop, interopercle; le, lateral ethmoid; me, mesethmoid; mpt, metapterygoid; mx, maxilla; op, opercle; orb, orbitosphenoid; pa, parietal; pas, parasphenoid; pc, posterior ceratohyal; pcl, postcleithrum; pl, palatine; pmx, premaxilla; pop, preopercle; pr, non-ossified projection of dentary; pts, pterosphenoid; q, quadrate; ra, retroarticular; scl, supracleithrum; soc, supraoccipital; sop, subopercle; sph, sphenotic; sr, sclerotic ring; str, cleithral strut; vhy, ventral hypohyal. straight, extending for most of the basicranial length. The vomer is not clearly visible. The mesethmoid is thin and elongated; it extends posteriorly and partially overlies the anterior portion of the frontals. The lateral ethmoids are large and fan-shaped, with a broad dorsal portion. The specimen MGPUP 11611 / 2 shows a complete and well-preserved infraorbital series (Fig. 4 A, B) comprising eight elements. The lachrymal is very large and irregular in shape. As in extant paralepidids, this bone is located horizontally on the snout, anterior to the orbit, and extends along the upper border of the maxilla (see Baldwin & Johnson, 1996); it is ornamented with a few radial grooves along its dorsal margin. The second and third infraorbitals are small and triangular. The posterior process of the posterior five infraorbitals is well developed with a lanceolate or subquadrangular outline. The antorbital is small, is irregular in shape and lies dorsal to the lachrymal. The nasal is subrectangular in outline and anteroposteriorly elongated. A unique subtrapezoid supraorbital is also recognizable in MGPUP 11611 / 2. The sclerotic ring is partially visible in MCSNV IG. 23601 / 2 (Fig. 3 A). The premaxilla is elongated, not protractile and closely attached to the maxilla throughout its length, thereby excluding it from the mouth gape (Figs 3 B, 4 A, B). It has a moderately developed and posteriorly directed ascending process with a wide ovoid premaxillary fenestra. Anteriorly, the premaxilla bears small and pointed retrorse teeth. The edentulous maxilla is thin, anteroposteriorly elongated, with a broad posterior margin. A single short supramaxilla closely associated with the posterodorsal margin of maxilla is well expanded in specimen MGPUP 1161 / 2 (Fig. 4 A, B). The dentary is slender with a nearly straight dorsal margin; three to four foramina of the laterosensory system are visible in the anterior portion of the dentary (Fig. 4 A, B). The lower jaw teeth are large, without a corrugate pattern or serrate margins. † Holosteus esocinus is characterized by a vertical, moderately developed symphysis, probably accommodated between the two contralateral premaxillae when the mouth was closed (see Harry, 1953 a). Anterior to it, there is a long, non-ossified and horizontally directed extension, projecting well beyond the anterior tip of the upper jaw (Fig. 3 A, B); a similar structure characterizes many derived alepisauroids (Harry, 1953 a, b), although it is extremely elongated in † H. esocinus with its length being contained about five to six times in head length. The anguloarticular is robust, triangular in shape and penetrates the dentary for about the half of its length. The retroarticular is small and curved. The palatine is robust and elongated; as in all the alepisauroid fishes, the palatine supports a remarkable series of well-developed pointed teeth (Fig. 3 A); the palatine teeth are long, without serrations or corrugate pattern and increase in size posteriorly in the series. As in other paralepidids (see Baldwin & Johnson, 1996), the palatine terminates anteriorly with a long process articulating with the premaxilla. The quadrate is fan-shaped, with a slight concavity along the dorsal margin; the quadrate – articular joint is located just below the posterior margin of the orbit, in a position that significantly differs from that characteristic of the Oligocene † P. mariae in which this articulation lies anterior to the orbit (see Prokofiev, 2005). The symplectic is not clearly recognizable. The metapterygoid is subtriangular in shape, with rounded margins; it does not extend anteriorly as in synodontids (Baldwin & Johnson, 1996), and its posterior margin articulates with the hyomandibula. The ectopterygoid is slender and gently curved. The endopterygoid is anteroposteriorly elongated. There is no trace of pterygoid teeth. The hyomandibula is short, robust and oriented vertically; it bears two dorsal articular heads and a distinct opercular process emerging dorsally from its posterior margin. The preopercle is large and crescent-shaped, with the vertical arm slightly longer than the ventral one; the corner formed by the junction of the two arms of the preopercle is ornamented with several radial striae. The opercle is the largest bone of the opercular series; it is subrectangular in outline, about 1.5 times broader than deep, with a robust condyle for the articulation with the hyomandibula. The subopercle is long, with a rounded posterior margin. The interopercle is long and slender. The hyoid apparatus is located just under the orbital region (Fig. 3 A, B). The anterior ceratohyal is slender, is subrectangular in shape, constricted in the middle and characterized by an indented ventral margin. The posterior ceratohyal is about the same length as the anterior ceratohyal and robust. There are eight branchiostegal rays, four of which articulate with the anterior ceratohyal; the first three branchiostegals are closely associated with each other along the anterior side of the indentation. The posterior two branchiostegals associated with the posterior ceratohyal are very close to each other and insert on the posteroventral corner of the bone. The ventral hypohyal is small and subquadrangular in shape. The dorsal hypohyal as well as the gill arches are not recognizable in the available specimens. The vertebral column consists of 106 – 112 vertebrae, of which 75 – 78 are abdominal and 31 – 34 are caudal; caudal vertebrae represent about 30 % of the total number. The vertebral centra are subquadrangular, slightly higher than long. Pre- and postzygapophyses are weakly developed throughout the entire vertebral column. Neural and haemal spines of the posterior 25 – 28 caudal vertebrae are thick and robust with a proximal broad laminar expansion, especially on the posteriormost 10 – 12 caudal vertebrae. There are about 75 pairs of well-ossified pleural ribs, the first of which seems to originate on the fourth vertebra. Intermuscular bones are well developed and extend from the abdominal to the caudal region. Epineurals originate laterally on the neural arches; it is unclear whether these are fused to the arches on the anterior vertebrae or not. The epineurals are forked proximally (most of them are bifid and in a few cases trifid; Fig. 5 A) from vertebrae 12 – 15 to the end of the abdominal region, thereby resembling the condition of certain primitive ipnopoids and chlorophthalmoids (see Patterson & Johnson, 1995; Baldwin & Johnson, 1996); most of the branches of the forked epineurals appear to be autogenous, unattached to the axial skeleton. Epineurals are long and thin, reaching the dorsal margin of the body; those of the caudal region are more robust and thick. Like in extant paralepidids, epipleurals seem to originate on the first vertebra, and at least the two anterior are autogenous (Fig. 4 A, B). † Holosteus esocinus shows an extended series of epipleurals associated with all the abdominal vertebrae and most of the caudal vertebrae, resembling the condition of Paralepis and Arctozenus (Baldwin & Johnson, 1996). The epipleurals of the abdominal region are thin and long, reaching the ventral margin of the body. All the epipleurals are attached to the axial skeleton and are not forked distally or proximally. As in extant paralepidids, there are no epicentrals in † H. esocinus. The caudal skeleton is consistent with that of most extant paralepidids (Fig. 6 A, B), although some unique features can be recognized. The first preural and first ural centra appear to be fused to each other. The second ural centrum is small and subtriangular in shape. There are six autogenous hypurals, of which the first and the second are fused into a single plate, resembling the condition observed in Arctozenus and Lestrolepis (see Fujita, 1990; Baldwin & Johnson, 1996). The autogenous parhypural is long and slender. Two autogenous uroneurals are recognizable, of which the anterior one is the largest. There are two thin and slender epurals. There are no urodermals. The caudal fin is large and externally symmetrical, with a nearly concave posterior margin; it contains 19 – 20 caudal-fin rays with segmentation beginning on the proximal half of each ray; the proximal portion of the principal caudal-fin rays is not modified in any way. The proximal ends of most of the caudal-fin rays of the upper lobe extensively overlap the associated hypurals, resulting in a sort of epaxial hypurostegy (Fig. 6 A, B). There are about 10 – 12 upper and 10 – 12 lower procurrent rays. There are two well-developed bony fulcral scales, nearly ovoid in shape located just in front of the upper and ventral caudal-fin lobes, resembling the condition characteristic of certain aulopoids and chlorophthalmoids (Sulak, 1977; Russell, 1999). The number and morphology of supraneurals is unclear. The dorsal and anal fins are displaced to the posterior half of the body, just in front of the caudal-fin (Figs 1, 2, 5 B). The dorsal-fin origin is located slightly behind that of the anal fin; the dorsal fin is triangular in shape and contains 18 – 20 distally segmented rays decreasing in size posteriorly. There is no trace of an adipose fin between the dorsal fin and caudal fin. The anal fin has a long base and contains 19 – 22 distally segmented rays; as in all alepisauroids, its outer margin is deeply indented anteriorly. Dorsal- and anal-fin rays are supported by a similar number of pterygiophores; the posterior pterygiophores of both the dorsal and anal fins do not appear to be fused or proximally modified. The post-temporal is not recognizable in the available material. The supracleithrum is spatulate, laminar and subtriangular in shape. The cleithrum is crescent-shaped with a broad ventral arm. It bears a distinctive paddle-shaped projection (= cleithral strut of Baldwin & Johnson, 1996) along its posterior margin; the presence of such a projection typically characterizes the most derived paralepidids (Davis, 2010; Fig. 3 A, B; Table 2). The number of postcleithra is unclear, but at least one is visible along the posterior margin of the supracleithrum; its position and size, as well as its subcircular shape, are reminiscent of the first postcleithrum of Paralepis atlantica and Stenomosudis rothschildi (see Rofen, 1966; Baldwin & Johnson, 1996). The coracoid is robust and large. The cleithrum – coracoid articulation is located near the anteroventral end of the cleithrum. At least three or four proximal radials can be recognized. The pectoral fin sits low on the body flanks and contains 16 – 18 rays, with their proximal portions (= spurs of Sato & Nakabo, 2002) being almost equal in size; the first pectoral-fin ray is the longest of the series. As in extant paralepidids, the pectoral-fin base seems to be horizontally oriented and inserts along the ventrolateral surface of the body. The pelvic fins are abdominal and displaced slightly behind the mid-length of the body, well anterior to the dorsal fin origin. There are 10 – 12 pelvic-fin rays. The basipterygia are elongated, joined to each other medially and without any trace of posterior or lateral pelvic processes. The body is totally naked without any evidence of body or lateral-line scales. Because of the lack of body squamation, the pigmented peritoneal membrane is clearly visible in well-preserved specimens (e. g. MCSNV IG. 23601 / 2). † Holosteus esocinus has a broad, uniform, mid-dorsal brown-pigmented band along the dorsum, extending from the head to the caudal fin, similar to the body pigmentation pattern of the naked genera like Lestidium (Harry, 1953 b). There is no evidence of pigmentation along the ventral surface of the body. There are no traces of preserved chromatophores.	en	Marramà, Giuseppe, Carnevale, Giorgio (2017): Morphology, relationships and palaeobiology of the Eocene barracudina † Holosteus esocinus (Aulopiformes: Paralepididae) from Monte Bolca, Italy. Zoological Journal of the Linnean Society 181 (1): 209-228, DOI: 10.1093/zoolinnean/zlw029, URL: http://academic.oup.com/zoolinnean/article/181/1/209/3782581/Morphology-relationships-and-palaeobiology-of-the
