A New Genus and Species of Lutjanid Fish (Perciformes) from the Eocene of Bolca, Northern Italy

Studi e ricerche sui giacimenti terziari di Bolca, XV - Miscellanea paleontologica, 12, 2014: 15 - 22

A new genus and species of lutjanid fish (Perciformes) from the Eocene of Bolca, Northern Italy

ALEXANDRE F. BANNIKOV* - ROBERTO ZORZIN** (*Borisyak Paleontological Institute of the Russian Academy of Sciences, Moscow) (**Museo Civico di Storia Naturale, Verona)

Abstract

A new genus and species of lutjanid fish (Perciformes, Percoidei),Lessinia horrenda, is described based on a single skeleton from the Eocene locality of Bolca in northern Italy. An additional specimen from the same locality is identified asLessinia sp. In comparison with other lutjanids from Bolca, the new genus has a smaller mouth and weaker dentition. The new taxon extends the list of Bolca snappers known to date to five or six species and four genera. Key words: Perciformes, Lutjanidae, Eocene, northern Italy, Bolca locality.

Riassunto

Un nuovo genere e specie di lutianide (Perciformes, Percoidei), Lessinia horrenda, è descritto sulla base di un unico scheletro proveniente dall’Eocene di Bolca (Italia settentrionale). Un ulteriore esemplare, della stessa località, è identificato come Lessinia sp. Dal confronto con altri lutianidi di Bolca, il nuovo genere ha una bocca più piccola e una dentatura più debole. Il nuovo taxon estende l’elenco dei dentici di Bolca noti fino ad oggi a cinque o sei specie e quattro generi. Parole chiave: Percifomi, Lutjanidae, Eocene, Italia settentrionale, Bolca.

Introduction by Bannikov (2006) and attributed to the new genus Goujetia. Finally, Bannikov (2006) showed that the spe- The fauna of the famous Monte Bolca locality, to the cies known as the serranid Serranus ventralis (Agassiz, north of Verona in northern Italy, is exceptionally rich 1833-1844; de Zigno, 1874) actually represents a snap- in marine Eocene fishes, especially of acanthoptery- per of the new genus Veranichthys, of which Serranus gians. However, in his systematic catalog of Bolca fish- rugosus Heckel, 1854 must be regarded as its junior syn- es Blot (1980) did not indicate any lutjanid fish from onym. In addition to the true lutjanids, there is also an there. Lutjanids, or snappers, are mostly reef-dwelling incertae sedis percoid in the Bolca fish fauna which is fishes presently widely distributed in tropical and sub- probably related to the Lutjanidae: Parapelates quindec- tropical marine waters. There are 17 genera and about imalis (Agassiz, 1836), as suggested by Bannikov (2008). 105 species of lutjanids in the Recent fauna (Allen, 1985; The field excavations executed in 2005-2011 by the Nelson, 2006). second listed author and his colleagues about 25 me- It has recenly been shown (Bannikov, 2006) that the ters higher up the steep hillside from the entrance to family Lutjanidae is represented in the Bolca fauna by the classic Pesciara cave site of the Monte Bolca local- at least four species in three genera (Ottaviania Sorbini, ity yielded many interesting materials, including new 1983, Goujetia Bannikov, 2006 and Veranichthys Ban- species of fishes and additional specimens of some fish nikov, 2006). Ottaviania mariae was briefly described taxa that had previously been known only on the ba- by Sorbini (1983) as a perciform fish of uncertain rela- sis of holotypes collected at the old Pesciara site during tionships, but it was subsequently redescribed by Ban- the nineteenth century. Among the materials collect- nikov and Zorzin (2004) in detail as a lutjanid per- ed at the new Pesciara site in 2007 is a single specimen haps related to the subfamily Etelinae. The species long of lutjanid fish that can be attributed to the new genus known as the sparid Dentex leptacanthus (Agassiz, 1833- and species described below. A poorly preserved speci- 1844; de Zigno, 1874) was shown by Bannikov (2006) men (IG 126422/126423) in the collection of the Museo to be a second species of Ottaviania. Similarly, Dentex Civico di Storia Naturale di Verona also belongs to this crassispinus Agassiz, 1839 was redescribed as a lutjanid new genus, but it probably represents a different spe- 16 ALEXANDRE F. BANNIKOV - ROBERTO ZORZIN cies; however, we prefer to wait for a better preserved Systematic Description specimen to be obtained before describing it. Thus, the list of the Bolca snappers known to date Order Perciformes can be extended to five or six species and four genera. Suborder Percoidei Even though they are relatively diverse taxonomically, Family Lutjanidae Gill, 1884 the lutjanids are not commonly collected at the Bol- ca locality; each of their species is represented to date Genus Lessinia gen. nov. by only one to a few specimens in the collections of the world’s museums. Type species Lessinia horrenda sp. nov., by monotypy and designa- Methods tion herein.

Some details of the specimens examined were best Diagnosis seen when the specimens were moistened with alcohol Dorsal profile of body somewhat more convex than during microscopic examination. ventral profile. Head relatively small. Maximum body Interneural and interhaemal spaces are numbered depth exceeds head length. Supraoccipital crest promi- based on the vertebra whose neural or haemal spine nent. Mouth small; lower jaw articulation under ante- forms the anterior border of the space, with the first rior half of orbit. Jaws with an outer row of small blunt space being between the first and second neural or hae- conical teeth and an inner band of minute sharp vill- mal spines (following Baldwin and Johnson, 1993; Ban- iform teeth. One or a few anterior premaxillary teeth nikov and Tyler, 1995; Tyler and Bannikov, 1997; etc.). canine-like. Preopercle finely serrated. 10+14 vertebrae. Abbreviations are as follows: Institutional: MCS- Hypurals 1-2 and 3-4 fused into two plates. Three strong NV – Museo Civico di Storia Naturale di Verona; An- supraneurals. Dorsal fin continuous, with 10 slender atomical: E – epural; H – hypural; HL – head length; spines and 11 soft rays. Anal fin with 3 graduated slender HPU – haemal spine of preural vertebra; Mx – maxil- spines and 8 rays. Pelvic fins inserted behind pectorals. la; NPU – neural spine of preural vertebra; Pal – pala- Outer pelvic-fin ray not elongated. Pectoral fins moder- tine; Ph – parhypural; Pmx – premaxilla; PU – preural ately long. Caudal fin deeply forked. Scales moderately vertebra; SL – standard length; U – ural vertebra; Un large and cycloid, each scale with numerous radii basal- – uroneural. ly. Lateral line complete, passes relatively low on flank.

Fig. 1 – Lessinia horrenda sp. nov., photograph of holotype MCSNV IGVR 71613, 152 mm SL; uppermost Lower Eocene, Monte Bolca locality in northern Italy, Pesciara site. Scale bar = 2 cm. A new genus and species of lutjanid fish (Perciformes) from the Eocene of Bolca, northern Italy 17

Fig. 2 – Lessinia horrenda sp. nov., reconstruction of entire skeleton based upon holotype MCSNV IGVR 71613; scales omitted.

Fig. 3 – Lessinia horrenda sp. nov., drawing of upper jaws of holo- Fig. 4 – Lessinia horrenda sp. nov., reconstruction of caudal skeleton type MCSNV IGVR 71613. Scale bar = 5 mm. based upon holotype MCSNV IGVR 71613; dark coloration marks outermost principal caudal-fin rays. Scale bar = 4 mm.

Etymology single plate; upper part of the Lower Eocene, Ypre- Named after the natural park Lessinia to the north of sian, zone Alveolina dainellii (Papazzoni and Trevisani, Verona; gender feminine. 2006); Monte Bolca locality, Pesciara cave site.

Referred species Referred specimens Type species and Lessinia sp. from the same locality. None.

Lessinia horrenda sp. nov. Etymology Figures 1 – 4 The species is named after the Latin word horrendus for marvelous, respectable. Diagnosis As for the genus. Description The body is moderately elongate, with a relatively short Holotype and rather slender caudal peduncle. The caudal pedun- MCSNV IGVR 71613; complete skeleton, 152 mm SL, cle depth is about 0.26 of the body depth. The head 18 ALEXANDRE F. BANNIKOV - ROBERTO ZORZIN is relatively small, its length (tip of snout to anterior with its posterior border widely notched. The branchial edge of upper part of pectoral girdle) almost 1.25 times bones and pharyngeal dentition are indistinct. less than the body depth. The head length is contained - Axial skeleton. There are 24 vertebrae, ten abdom- about 3.28 times in SL. The dorsal profile of the body inal and fourteen caudal, including the urostyle. The is slightly more convex than the ventral profile of the axis of the vertebral column is almost straight. Most of body. the vertebral centra are either square or slightly elon- - Head. The head is relatively deep, with its depth gate anteroposteriorly and constricted in the middle. equals to its length. The orbit is moderate and situat- The length of the caudal portion of the vertebral col- ed in the upper half of the head. The horizontal diam- umn is 1.6 times greater than the length of the abdom- eter of the orbit is 26-27% HL. The snout is pointed inal portion of the vertebral column. The neural spines and relatively short; its length is about 30% HL. The of the abdominal vertebrae increase in length posterior- mouth is relatively small and terminal; the lower jaw ly in the series; those of the first five vertebrae are some- articulation is situated approximately under the ante- what thicker than the others. The neural and haemal rior half of the orbit. The infraorbital bones are bad- spines of the caudal vertebrae are moderately slender ly damaged, but the lachrymal seems to be relatively and straight or slightly curved, and decrease in length deep. The neurocranium is relatively deep, with a high posteriorly in the series. The first haemal spine is slight- supraoccipital crest that is prominent and rounded an- ly expanded in the middle. The neural spines of the terodorsally. The ethmoid region is short. The narrow caudal vertebrae are similar to the corresponding hae- lateral ethmoid is prominent laterally, forming the an- mal spines in length and in the angle of posterior incli- terior border of the orbit. The parasphenoid is robust nation. Parapophyses were definitely present on at least and almost straight; it extends along the lower border of the six posteriormost abdominal vertebrae, but most of the orbit. The premaxilla has a well-developed long and them are not preserved. Pleural ribs are present on the slender ascending process and a massive articular pro- third to tenth vertebrae, these are relatively long and cess. A high triangular postmaxillary process is present slender. Epineurals are situated on a level of the ver- on the distal half of the premaxilla (Fig. 3). The alveolar tebral column, and most of them are obscured by the ramus of the premaxilla bears an outer row of relatively centra. small blunt conical teeth, and an inner band of minute - Pectoral girdle. Extrascapulars are absent. A post- sharp conical teeth. One anterior tooth is canine-like. temporal and supracleithrum are only partly recogniz- The maxilla is relatively short and expanded towards able. The cleithrum is an elongate, strong bone, and its rounded distal end. The maxillary head is crescent- somewhat S-shaped. The upper part of the cleithrum is shaped. There are no supramaxillae. The mandible is curved forward and situated below the third vertebra. relatively short and rather deep. The length of the man- The ventral postcleithrum is only moderately long and dible is about 39% HL. The dentary is notched poste- wedge-like in shape; it is directed posteroventrally. The riorly. The retroarticular process of the angulo-articu- coracoid is moderately narrow; there is a gape between lar is massive and deep. The dentary dentition is similar the lowermost portions of the cleithrum and coracoid. to that of the premaxilla except that an anterior canine- The scapula and pectoral radials are scarcely recogniza- like tooth is not evident. The hyomandibular shaft is ble. The pectoral fin is mostly hidden by scales and ribs; significantly inclined anteroventrally. The quadrate is however, it is evident that the fin is moderately long. subtriangular and moderately large. The symplectic The total complement of pectoral-fin rays is unknown. seems to be a simple strutlike bone. The ectopterygoid The pectoral fin is inserted under the fifth vertebra just evidently lacks a posterior extension separating the en- above the midpoint between the vertebral column and dopterygoid from the quadrate. The limits of the endo- the ventral profile of the body. and metapterygoid are unclear. The opercular region is - Pelvic fin and girdle. The pelvic bones are relative- moderately wide. The preopercle is only slightly curved ly strong and wedge-like. The anterior end of the basip- and moderately wide; its posterior edge is finely serrat- terygium is oriented anterodorsally and attaches rela- ed. The opercle is wide and thickened along its anterior tively high on the cleithrum. The pelvic fin has a spine border. There are seven sabre-like branchiostegal rays; and five soft branched rays; it is inserted behind the the branchiostegals become longer and wider posteri- pectoral-fin base. The pelvic fin is moderately long. orly in the series. The hypohyals are much fragmented, - Supraneurals and dorsal fin. There are three very and the ceratohyal is exposed only anteroventrally; it strong supraneurals (predorsal bones); the first supra- seems to lack a beryciform foramen. The urohyal seems neural bears a long apical process directed anteriorly, to have a shape typical for lutjanids (Kusaka, 1974), whereas the third supraneural bears a long apical pro- A new genus and species of lutjanid fish (Perciformes) from the Eocene of Bolca, northern Italy 19 cess directed posteriorly. The predorsal formula (Ahl- imal segments. The interneural spaces below the dorsal strom et al., 1976) is /0+0/0+2/1+1/. Because in extant fin have the ventral shafts of one (usually) or two (the lutjanids the first supraneural aways precedes the first third, fifth, 11th, and 14th spaces) pterygiophores present. neural spine (Johnson, 1980, 1984), one can assume that - Anal fin. The anal fin originates under the third there was a slight post-mortem posterior displacement and ends under the ninth caudal vertebrae. There are of the lower end of the first supraneural. The dorsal fin three spines and eight soft segmented rays in the anal is relatively long-based and continuous; it originates fin; all of the soft rays are branched. The spines are slen- over the fourth abdominal vertebra and terminates der and very closely spaced; the third spine is at least over the ninth caudal vertebra. There are ten dorsal-fin 1.2 times longer than the second spine and 2.3 times spines and eleven soft segmented rays; the last ray is not longer than the first spine. The first two spines are su- produced. The dorsal-fin spines are slender. The first pernumerary. The longest anal-fin soft ray is not long- two spines are supernumerary; these are closely spaced er than the third anal-fin spine, which is slightly short- and shorter than the third to tenth spines, which are er than the longest dorsal-fin spine. The first anal-fin more or less equal in length. The first spine is 1.7 times pterygiophore is moderately long, sturdy, and wedge- shorter than the third. The first dorsal-fin soft ray is like in shape; it is inclined at an angle of about 55° to unbranched, whereas all the others are branched. The the body axis. The succeeding anal-fin pterygiophores longest soft ray of the dorsal fin is somewhat longer are slender and relatively strongly inclined; these gradu- than the last dorsal-fin spine. The length of the base ally decrease in length posteriorly in the series. The last of the soft portion of the dorsal fin is 1.17 times long- anal-fin pterygiophore is almost horizontally oriented. er than the base length of the spiny portion of the dor- The six posteriormost pterygiophores are trisegmental. sal fin. There are a total of 19 dorsal-fin pterygiophores. Most of the anal-fin pterygiophores are situated below The ventral shafts of the two anterior dorsal-fin ptery- the tips of the haemal spines, and thus they do not en- giophores are only slightly inclined, whereas posteriorly ter the interhaemal spaces. in the series the pterygiophores become more strongly - Caudal fin and skeleton. The terminal centrum is inclined, and the posteriormost pterygiophores are al- composed of the fusion of PU1, U1 and U2 (Fig. 4). most horizontally oriented. The pterygiophores of the The neural spine of PU2 is a short crest. The fifth hy- dorsal-fin spines are expanded anteroposteriorly, and pural, parhypural, and haemal spines of PU2 and PU3 each bears a longitudinal strengthening ridge. The first are autogenous. Hypurals 1-2 and 3-4 are fused into two pterygiophore is the largest; the succeeding pterygio- plates. There are three epurals which decrease in length phores become gradually shorter and narrower. At least posteriorly in the series. There is a small hypural diaste- the five posteriormost pterygiophores are trisegmen- ma (Monod, 1968). There are two uroneurals (the first tal, with their medial segments unfused with the prox- as a stegural). The neural and haemal spines of PU3 are

Fig. 5 – Lessinia sp., photograph of specimen MCSNV IG 126423, 179 mm SL; uppermost Lower Eocene, Monte Bolca locality in northern It- aly, Pesciara site. Scale bar = 2 cm. 20 ALEXANDRE F. BANNIKOV - ROBERTO ZORZIN longer and stouter than those of the preceding verte- the soft portion, whereas in the holotype of L. horrenda bra. The caudal fin is large and forked; it has 17 princi- sp. nov. the base length of the soft portion of the dor- pal rays, 15 of which are branched (I,8-7,I). The upper sal fin is longer. It is unlikely that these differences are lobe of the caudal fin perhaps is slightly longer than the caused by differing states of preservation, and it is dif- lower lobe. There are about 10 dorsal procurrent rays. ficult to explain them by possible sexual dimorphism; The number of ventral procurrent rays is less precise- most likely the two specimens under consideration be- ly known because the several anteriormost elements are long to two different species of the same genus. MCS- not preserved. NV IG 126422/126423 is somewhat less well preserved - Squamation. The scales are cycloid and moderately than the holotype of L. horrenda sp. nov., and some de- large; each scale bears numerous radii in the basal field. tails of its structure are not clear. Therefore, until a bet- The number of radii varies from six (breast scales) to 11. ter preserved specimen becomes available, we prefer to The scales cover the body and cheek. The lateral line is regard MCSNV IG 126422/126423 as Lessinia sp. very slightly arched and low in position; it descends to the level of the vertebral column near the 17th vertebra. Some scales bear minute spots of dark pigment. Discussion - Measurements. The following measurements are presented as a percent of SL (152 mm) of the holotype, We attribute Lessinia gen. nov. quite confidently to the to the nearest half mm: percoid family Lutjanidae. The new genus has a charac- Head length from tip of snout to posterior border of teristic lutjanoid overall appearance, with a single dor- opercle: 30.5 sal fin, terminal mouth, high position of the orbit, sev- Maximum body depth: 38 en branchiostegal rays, and supramaxilla absent, as well Depth of caudal peduncle: 10 as the dentition typical for lutjanoids (Johnson, 1980): Distance between tip of snout and spiny dorsal fin: 37 jaws with a canine, an outer row of larger conical teeth, Distance between tip of snout and soft dorsal fin: 59.5 and an inner band of smaller conical teeth. The dorsal Distance between tip of snout and anal fin: 64 (X, 11) and anal fin (III, 8) counts of Lessinia both are Distance between pelvic fin and anal fin: 21 within the range recorded in the Lutjanidae. Length of base of spiny dorsal fin: 22 In his revision of extant lutjanoids, Johnson (1980) Length of base of soft dorsal fin: 25.5 recognized the superfamily Lutjanoidea consisting of Length of base of entire dorsal fin: 50 two families, Lutjanidae and Caesionidae. The Lutja- Length of base of anal fin: 21 nidae was subdivided by Johnson into four subfami- Length of first spine of dorsal fin: 7.5 lies, Etelinae, Apsilinae, Paradicichthyinae and Lutjani- Length of longest spine of dorsal fin: 13 nae. Johnson’s classification was accepted by succeeding Length of longest soft ray of dorsal fin: 14 authors (e.g., Allen, 1985; Carpenter, 1990), but sub- Length of third spine of anal fin: 10.5 sequently Johnson (1993) recommended treating cae- Length of longest soft ray of anal fin: 10 sionines as a subfamily of the Lutjanidae, which was Length of longest ray of pelvic fin: 17.5 originally accepted by Nelson (1994). Later, Nelson Length of longest ray of caudal fin: ca. 35 (2006) changed back to treating the Caesionidae as a Preorbital distance: 9 separate family. Lessinia gen. nov. is easily distinguish- Horizontal diameter of orbit: 8.5 able from caesionines by its different dorsal-fin formu- Length of lower jaw: 12 la, absence of the separate ossification of the premaxillary ascending process that is characteristic for the Remarks caesionines, and a broad-based rather than finger-like Specimen MCSNV IG 126422/126423 (Fig. 5) is simi- postmaxillary process. lar in many respects to the holotype of Lessinia horren- Among the morphological characters of Lessinia da sp. nov., and has to be attributed to the same genus. gen. nov., two have not previously been reported for However, in certain morphological character states it the Recent lutjanids (Johnson, 1980, 1984): the predor- is different from L. horrenda sp. nov., having a some- sal configuration and cycloid scales. The first supraneu- what more elongate body, lower supraoccipital crest, ral of extant snappers precedes the first neural spine, longer ethmoid region, several canine-like teeth ante- whereas it enters the first interneural space in L. horren- riorly on the premaxilla (vs. a single tooth in L. horrenda sp. nov. (this character state is less clear in Lessinia da sp. nov.). Also, MCSNV IG 126422/126423 has the sp.). However, as mentioned above, one can assume a spiny portion of the dorsal fin longer at the base than slight post-mortem posterior displacement of the low- A new genus and species of lutjanid fish (Perciformes) from the Eocene of Bolca, northern Italy 21 er end of the first supraneural in the holotype of the that the incertae sedis Oligocene percoid fish genus Oli- new species. Concerning the cycloid scales, such scales ganodon Bannikov from the Caucasus and Central Eu- have already been reported for one of the two species rope (Bannikov, 2010) may actually be attributable to of the Eocene lutjanid genus Ottaviania, and for the the Lutjanidae. However, it is from Bolca that the fossil genus Veranichthys, also from Bolca (Bannikov, 2006). Lutjanidae are by far the best known. The family-level scale characteristic certainly has slightly changed throughout the last 50 millions of years of lutjanid history. Acknowledgements In its dorsal and anal fin counts, deeply forked caudal fin and fused hypurals 1-2 and 3-4, Lessinia gen. nov. We are very grateful to Dr. Giuseppe Minciotti, di- is closer to the lutjanid subfamilies Etelinae and Apsil- rector of the Museo Civico di Storia Naturale di Ve- inae than to the Paradicichthyinae and Lutjaninae (see rona, for providing funding for the travel of AFB in Johnson, 1980). But the Eocene genus more closely re- 2013 to Verona to engage in this and other studies. We sembles apsilines rather than etelines in a number of its also thank Dr. Anna Vaccari, collection manager, for character states. Unlike the Etelinae, Lessinia gen. nov. her assistance, and Mrs. Bruna Burato for her aid with has the ultimate ray of the dorsal and anal fins equal to the literature on Bolca fishes. We are much indebted or shorter than the penultimate, and a relatively high su- to Mr. Francesco Sorbini (Verona) for the photographs praoccipital crest (vs. low crest in the Etelinae). Lessin- and Dr. James C. Tyler of the Smithsonian Institution ia gen. nov. differs from the Apsilinae in having some- for reviewing and correcting the English in the manu- what shorter pectoral fins and slightly more numerous script. The work of AFB was supported by the Russian dorsal-fin soft rays (11 vs. 9 or 10 in the Apsilinae). The Foundation for Basic Research, grant no. 14-04-00005. predorsal configuration of Lessinia horrenda sp. nov. (/0+0/0+2/1+1/) is different from those of both the Etelinae (0/0+0/2/1+1/) and Apsilinae (0/0/0+2/1+1/). Literature Cited But the Lessinia condition could be more easily trans- formed into the predorsal configuration of the Apsili- AGASSIZ L., 1833-1844. Recherches sur les poissons nae than into that of the Etelinae, just by the simple an- fossiles. Petitpierre, Neuchâtel: Tome IV: 296 pp. terior shift of the first supraneural. Such character states AHLSTROM E.H., BUTLER J.L., SUMIDA B.Y., of Lessinia gen. nov. as its relatively small head, relative- 1976. Pelagic stromateoid fishes (Pisces, Perciformes) of ly deep body and small mouth which differentiate this the eastern Pacific: kinds, distributions and early life his- genus from the other Eocene lutjanids, have their an- tories and observations on five of these from the north- alogues in some apsilines, especially in the genus Para- west Atlantic. Bulletin of Marine Science, 26: 285-402. caesio Bleeker, 1875, some species of which also have a ALLEN G.R., 1985. FAO species catalogue. Vol. 6. deeply forked caudal fin. The dentition of the new ge- Snappers of the world. An annotated and illustrated nus is much weaker than in the Eocene genera Ottavi- catalogue of lutjanid species known to date. FAO Fish- ania Sorbini, 1983, Goujetia Bannikov, 2006 and Veran- eries Synopsis, (125) Vol. 6: 208 pp. ichthys Bannikov, 2006 from Bolca, resembling that of BALDWIN C.C., JOHNSON G.D., 1993. Phylog- the modern apsiline genus Apsilus Valenciennes, 1830 eny of the Epinephelinae (Teleostei: Serranidae). Bulle- (Allen, 1985). Another extant apsiline genus, Paraprist- tin of Marine Science, 52 (1): 240-283. ipomoides Kami, 1973, has dentition even weaker than BANNIKOV A.F., 2006. Fishes from the Eocene of in Lessinia gen. nov., because it has no canine-like teeth Bolca, northern Italy, previously classified in the Spari- present (Johnson, 1980; Allen, 1985). dae, Serranidae and Haemulidae (Perciformes). Geodi- Thus, Lessinia gen. nov. represents a distinct genus versitas, 28 (2): 249-275. of the Lutjanidae. It seems to be closer to the Apsilinae BANNIKOV A.F., 2008. Redescription of “Pelates” than to the other subfamilies of lutjanids and can be quindecimalis Agassiz, 1836, percoid fish (Perciformes) tentatively placed in that subfamily. from the Eocene of Bolca, Italy. Miscellanea paleontolog- Other than in the Eocene of Monte Bolca where ica n. 9. Studi e Ricerche sui Giacimenti Terziari di Bol- at least four of their genera are present, snappers are ca, 12: 57-64. scarcely known at all in the fossil record. Patterson BANNIKOV A.F., 2010. Fossil Vertebrates of Russia (1993) indicated that the late Eocene Hypsocephalus at- and Adjacent Countries. Fossil Acanthopterygians Fishes lanticus Swift et Ellwood from Florida was the oldest lu- (Teleostei, Acanthopterygii). GEOS, Moscow: LXI+244 tjanid based on osteological material; and it is possible pp. [In Russian]. 22 ALEXANDRE F. BANNIKOV - ROBERTO ZORZIN

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