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Research article

um:lsid:zoobank.org:pub:0F037EA5-48E4-4029-9DCA-B04CC2366A53

Gladiopycnodontidae, a new family of pycnodontiform fishes from the

Late Cretaceous of Lebanon, with the description of three genera

Louis TAVERNE' & Luigi CAPASSO^

^ Royal Belgian Institute of Natural Seienees, Department of Paleontology, Vautierstreet, 29,

B-1000 Brussels, Belgium. E-mail: [email protected]

^Museo Universitario delEUniversita “G. d’Annunzio” di Chieti-Peseara, Piazza Trento e Trieste 1,

1-66100 Chieti, Italy. E-mail: [email protected]

i um:lsid:zoobank.org:author:0CF81641-lDDl-4CBD-9735-FlFE7EB0BCF5

" urn:lsid:zoobank.org:author:C79C14CF-ClD7-48E3-9BA9-CAD8AA0909F7

Abstract. The osteology of Gladiopycnodus karami gen. et sp. nov., of Monocerichthys scheuchzeri

gen. et sp. nov. and of Rostropycnodus gayeti gen. et sp. nov., three new fossil fishes from the marine

Cenomanian (Eate Cretaeeous) of Eebanon, is studied in detail. Some of their eranial eharaeters

and the presenee of a posteoelomie bone elearly refer these fishes to the order Pyenodontiformes.

However, they differ from all other deseribed Pyenodontiformes by two important eharaeters. Their

snout is elongated as a rostrum, formed by the enlarged prefrontal and the toothless premaxilla, with

this premaxilla sutured by its upper margin to the lower margin of the prefrontal. Their peetoral

fin is replaeed by a strong spine artieulated with the eleithrum. These two apomorphies justify the

ereetion of a new family, the Gladiopyenodontidae. The skull of Monocerichthys scheuchzeri

sp. nov. does not differ greatly from a elassieal pyenodontiform skull and this speeies seems to be the more

primitive member of this new family. Gladiopycnodus karami gen. et sp. nov. and Rostropycnodus gayeti

gen. et sp. nov. are muehmore speeialized. They share some apomorphies not present in Monocerichthys

scheuchzeri gen. et sp. nov., i. e., an extremely long rostrum and an elongated first anal pterygiophore

that sustains with the posteoelomie bone a strong and long anal spine. Gladiopyenodontidae fam.

nov. and Coeeodontidae share a series of apomorphies that justify the ereetion of a new superfamily,

Coeeodontoidea, grouping these two families.

Key words. Pyenodontiformes, Coeeodontoidea superfam. nov., Gladiopyenodontidae fam. nov.,

Gladiopycnodus karami gen. et sp. nov., Monocerichthys scheuchzerii gen. et sp. nov., Rostropycnodus

gayeti gen. et sp. nov., osteology, relationships, Cenomanian, Eebanon.

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/eit.2013.57

Introduction

Pycnodontiform fishes are by far the largest marine halecostome fossil lineage, that range in time from

the Eate Triassic to the Middle Eocene and have an almost worldwide distribution (Nursall 1996a;

European Journal of Taxonomy 57 : 1-30 ( 2013 )

Kriwet 2001). They are often found in marine and more rarely in braekish or freshwater deposits. Most

of them possess a deep and laterally eompressed body. They generally have a durophagous mode of

feeding thanks to their molariform teeth on the vomer and the preartieulars (Nursall 1996a: fig. 3). Some

speeialists eonsider them as elosely related to the teleosts (e.g., Nursall 2010).

For a very long time, all pyenodontiform fishes were ranged in the order Pyenodontiformes (e.g., Nursall

1996b; Poyato-Ariza & Wenz 2002; Kriwet 2005). But reeently, a new superorder Pyenodontomorpha

was ereetedto aeeomodate them, eomprisingtwo orders: the Gyrodontiformes for the families Mesturidae

and Gyrodontidae and the Pyenodontiformes (new usage for the former Pyenodontoidei) for the families

Brembodontidae, Coeeodontidae, Trewavasiidae and Pyenodontidae (Nursall 2010). However, some

speeialists disagree with sueh a division and eonsider the Gyrodontiformes as polyphyletie (Poyato-

Ariza pers. eom., April 2013).

Pyenodontiform fishes are well known in the marine Late Cretaeeous of Lebanon (e.g., Pietet 1850;

Davis 1887, 1890; Hay 1903; Poyato-Ariza & Wenz 2002, 2005; Forey et al. 2003; Capasso et al.

2009; Gayet et al. 2012) and they inelude a few highly speeialized families, sueh as Gebrayeliehthyidae

(Nursall & Capasso 2004) and Coeeodontidae [ineluding Trewavasiidae] (Gayet 1984; Kriwet 2004;

Nursall & Capasso 2008; Capasso et al. 2010).

Capasso’s paleontologieal eolleetion in Chieti (Italy) eontains numerous still undeseribed pyenodontiform

fishes from Lebanon. Some of them represent a new remarkable family eontaining about ten genera.

The fishes belonging to these genera are not deep-bodied, but fusiform in shape, exeept for one very

speeialized genus with a rounded body. Their snout is more or less elongated, forming a rostrum that

anteriorly protrudes above the lower jaw. The anterior extremity of this rostrum is eonstituted by the

enlarged prefrontal only and not by the premaxilla that usually assumes this role in a pyenodontiform

snout. The long, broad, toothless premaxilla is loeated below the prefrontal and is sutured along its upper

margin to this prefrontal and so strengthens the rostrum. Their peetoral fin is lost and replaeed by a strong

spine. Some speeies of the new family possess frontal, oeeipital, nuehal and eleithral horns or spines. The

Cenomanian Lebanese speeies ''Coccodus’' lindstroemi Davis, 1890 (not a true Coccodus Pietet, 1850,

see Poyato-Ariza & Wenz 2002: 145) belongs to this new lineage (Taveme & Capasso, work in prep.).

The aim of our paper is to deseribe three members of this new family, the less speeialized and the two most

speeialized genera. We will also diseuss the systematie position of the family within pyenodontiform

fishes. Other papers are in preparation and will eomplete the study of the group.

Material and methods

The speeimens herein studied belong to the Luigi Capasso eolleetion (CLC) in Chieti (Italy), that is

legally registered by a deeree of the Ministero per i Beni e le Attivita Culturali, date of Oetober IP*’,

1999, following the disposition of the Italian law 1089/39. The Soprintendenza per i Beni Areheologiei

delFAbruzzo - Chieti has authorized the authors to study this eolleetion by a letter bearing the date of

May 5*h, 2011 (referenee: MBAC-SBA-ABR PROT 0004537 05/05/2011 Cl. 34.25.01/2.1).

The speeimens were studied with a Leiea-Wild M 8 stereo mieroseope. The figures were drawn by the

first author (LT). Aspersions with ethanol were used to improve the observations.

List of abbreviations used in text-figures

angular

ART

artieular

ASPH

autosphenotie

CLT

eleithrum

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

DHYOM

dermohyomandibula

dentary

DPTE

dermopterotie

DSOC

dermosupraoeeipital

DSPH

dermosphenotie

ENPT

entopterygoid (= endopterygoid)

EPCO 1-6

epiehordals 1 to 6

frontal

HAEM

haemal areh

HAEMEP

haemal spine

HCET

hypereleithrum (= supraeleithmm)

HYCO 1-12

hypoehordals 1 to 12

HYOM

hyomandibula

lORB 1-5

infraorbitals 1 to 5

EEP

lepidotriehium

METH

mesethmoid

MPT

metapterygoid

maxilla

NEUR

neural areh

NEUREP

neural spine

nuehal horn

operele

OSPH

orbitosphenoid

parietal

POET

posteleithrum

PCOEE

posteoelomie bone

PEEV

pelvie bones

PMX

premaxilla

POP

preoperele

PRART

preartieular

PRFR

prefrontal (= laterodermethmoid ?)

parasphenoid

posttemporal

quadrate

RAD

pterygiophores

RADd. 1-13

dorsal pterygiophores (= radials) 1 to 13

RAD a

anal pterygiophores

RADI

first anal pterygiophore

SCb

body seale

SCe

eaudal seale

SCE

selerotie bone

SOU d.

dorsal ridge seute

SCUp.

pelvie seute

SOU po.

posteloaeal seute

SPl a.

spine of the anal fin

SPlp.

peetoral spine

supratemporal

sympleetie

vomer

ol. f

olfaetive fossa

European Journal of Taxonomy 57 : 1-30 ( 2013 )

Results

Subclass Actinopterygii Klein, 1885

Series Neopterygii Regan, 1923

Division Haleeostomi Regan, 1923 Patterson 1973

Superorder Pyenodontomorpha Nursall, 2010

Order Pyenodontiformes Berg, 1937 sensu Nursall 2010

Superfamily Coeeodontoidea superfam. nov.

um:lsid:zoobank.org:aet:F0BC595E-F862-4373-860B-CB749BlEC5E4

Diagnosis

Pyenodontiformes primitively with a fusiform body, devoid of dorsal and ventral apex. Skull bearing

horns or spines. Dermohyomandibula present. Preoperele enlarged and mueh larger than the exposed part

of the hyomandibula-dermohyomandibula. Small operele present. Infraorbitals eovering only a small

part of the eheek. Earge peetoral girdle elosely assoeiated to the skull, forming a sort of eephalo-thorax.

Earge supratemporal artieulated with the dermosupraoeeipital, the parietal and the dennopterotie. Earge

posttemporal sutured to the supratemporal. Coronoid proeess of the preartieular weakly developed.

Enlarged hypereleithrum. Cleithrum greatly hypertrophied, with a ventral posterior proeess. Eess than 20

vertebral segments before the epiehordal series. Neural and haemal spines short. Dorsal fin primitively

short and loeated on the middle of the baek. Anal fin primitively short and largely separated from the tail.

East hypoehordal elements enlarged but not hypertrophied. One or no posteloaeal seute.

Family Gladiopyenodontidae fam. nov.

um:lsid:zoobank.org:aet:66FQ4581-AFC7-47B5-A6E8-7FBQ935ADD81

Diagnosis

Small Coeeodontoidea superfam. nov.. Snout elongated and forming a rostrum outpaeing anteriorly the

lower jaw level. Anterior tip of the rostrum formed by the long and broad prefrontal only. Premaxilla

long, broad, toothless and sutured all along its dorsal margin with the prefrontal. Peetoral fin lost and

replaeed by a strong spine artieulated with the eleithrum. Body entirely eovered with small fiake-like

seales or with seutes.

Genus Gladiopycnodus gen. nov.

um:lsid:zoobank.org:aet:623891D8-C2AE-4814-91C3-CBE48D098B88

Type-species

Gladiopycnodus karami gen. et sp. nov. (by monotypy)

Diagnosis

As for the speeies (monospeeifie genus).

Etymology

From the Eatin gladius, sword, referring to the shape of the peetoral and anal spines. The generie name

Pycnodus is added.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Gladiopycnodus karami gen. et sp. nov.

Figs 1-6

um:lsid:zoobank.org:aet:BE468D57-6344-4EA5-8F40-2AQ646799E2C

Diagnosis

Gladiopyenodontid with an extremely elongated snout, forming a sword-like rostrum horizontally

oriented and greatly outpaeing the lower jaw level. Anterior tip of the prefrontal ending in an aeuminate

point. Frontal long and narrow. Dermosphenotie and dermopterotie partly fused. Oral border of

premaxilla and maxilla toothless but bearing very small spines. Five infraorbitals, the first one larger

than the others. Earge lower jaw horizontally oriented. Earge preartieular bearing molarifonn teeth.

Preoperele greatly hypertrophied, with an elongated and dentieulated lower margin forming the ventral

border of the skull behind the lower jaw. Cleithrum vertieally oriented. Strong peetoral spine. Redueed

pelvie girdle present. Nuehal horn absent. Eong and robust posteoelomie bone. Dorsal fin with short

isolated rays forming a series of finlets. Anal fin beginning with an extremely long and strong spine that

greatly outpaees the tail level. Only a few short soft anal rays. Caudal fin with a eonvex posterior border.

Earge paired dorsal seutes between the skull and the tail. Body eompletely eovered with small flake¬

like seales irregularly imbrieated. Earge rounded seales on the eaudal pedunele. Two pelvie seutes. One

posteloaeal seute. No seute on the ventral margin of the body in the eaudal region.

Etymology

The speeies name of the new Eebanese fossil flsh is dedieated to Youssef Bey Karam (1823-1889), the

famous hero of Eebanon who lived during the time this eountry was ruled by the Ottoman Empire.

Fig. 1. Gladiopycnodus karami gen. et sp. nov. Holotype CEC S-393.

Fig. 2. Gladiopycnodus karami gen. et sp. nov. Reeonstruetion of holotype CEC S-393.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

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 meset hm oid 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 fiat, 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 comer 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 bmsh-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

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

sueh a fenestra does not exist in this speeies. The dermopterotie and the dermosphenotie are smaller

bones than the parietal. They are fused together dorsally but remain separated ventrally.

The parasphenoid is long and toothless. Avery small orbitosphenoid is present just behind the mesethmoid.

The other endoeranial bones of the brainease are hidden by the preoperele and the hyomandibula-

dermohyomandibula.

The upper jaw is a part of the rostrum and, thus, is loeated before the lower jaw, whieh does not

partieipate in the rostrum. The premaxilla is a very long and rather broad bone that is loeated 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 mueh shorter bone shaped as a lanee 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, redueed to its ventral

braneh and bearing two ineisiform prehensile teeth. The upper part of the anterior margin of the dentary

is ornamented with a few very small spines. The preartieular is the largest eomponent of the lower jaw.

Its inner faee is never visible, neither on the holotype nor on the other speeimens. However, some large

molariform teeth are visible under a broken part of the preartieular on the photo of one speeimen. The

angular is as deep as long. The artieular is longer but narrow.

Both the quadrate and the sympleetie artieulate with the lower jaw. As usual in pyenodontiform fishes,

the quadrate does not possess an ossified quadratie proeess. The sympleetie is a very robust eurved 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 eetopterygoid is visible.

There are five infraorbitals and a selerotie 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

speeimens. However, this bone eovers only a very small part of the eheek. The four other infraorbitals are

well developed but narrower than the first one and more or less redueed to their neurodermie eomponent.

DSOC

HYOM+

DHYOM

Fig. 3. Gladiopycnodus karami gen. et sp. nov. Reeonstruetion of the skull of holotype CEC S-393.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

The greatly hypertrophied preoperele 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

preoperele reaehes the dermosphenotie and the autosphenotie and exhibits a posterior hollow in whieh

the hyomandibula-dermohyomandibula fits. The operele is well developed but a lot smaller than the

preoperele. It is an ovoid bone with its dorsal and ventral extremities aeuminate. The branehiostegal

rays are not visible.

The hyomandibula and dermohyomandibula are intimately fused in a pyriform-shaped bone of the same

size as the operele and thus mueh smaller than the preoperele. Within this double eomposedbone, the small

hyomandibula is anteriorly loeated and has a smooth surfaee, whereas the larger dermohyomandibula is

posteriorly loeated and has a granulated surfaee. It is possible that a ventral braneh of the hyomandibula

is hidden by the preoperele. The hyoid bar is not visible.

2. The girdles (Figs 2, 4-5)

The peetoral girdle is pressed against the skull. The dermal bones of the peetoral girdle are ornamented

with small tubereles as those on the skull. The eleithrum is by far the largest element of the girdle. It is

a very deep, broad and vertieally oriented bone. There is a shortening in its upper part that ereates a sort

of hollow in its anterior margin. The operele fits in this hollow. The posttemporal is a very small and

more or less rounded bone wedged between the parietal, the supraoeeipital and the first dorsal seute.

The hypereleithrum (= supraeleithrum) is larger and amphora-like in shape. The peetoral fin is lost and

replaeed by a big pointed spine, artieulated on the eleithrum.

Fig. 4. Gladiopycnodus karami gen. et sp. nov. Reeonstruetion of the peetoral girdle and of the anal fin

of holotype CLC S-393.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

The pelvie girdle is hidden on the holotype beeause the fossilisation has pressed the peetoral and anal

spines against eaeh other. However, on one speeimen seen in photo, two small, obliquely oriented pelvie

bones are elearly visible between these two spines and some fragments of short pelvie rays appear under

the pelvie seutes.

3. The axial skeleton

The body is fusiform. It is diffieult to deseribe the axial skeleton in a detailed way, as it is generally more

or less eovered by the seales and it is not possible to eount the vertebral segments. The holotype shows a

few fragments of neural and haemal spines. On one of the speeimens seen on photo, many seales on the

body are lost and we ean observe that the notoehord is nearly eompletely surrounded by the neural and

haemal arehes. The neural and haemal spines are well developed but rather short. Eaeh of them bears a

broad anterior wing. The spines are not interloeked together by interdigitating sutures.

The presenee of ribs is uneertain, the situs viscerum being hidden by the gigantie preoperele, the operele

and the peetoral girdle.

The post-eoelomie bone is long and very robust. It reaehes the vertebral axis dorsally and the inferior

border of the fish ventrally. Its broad ventral part is baekwardly eurved.

4. The dorsal and anal fins (Figs 2, 4)

The dorsal fin is eomposed of about ten very short, branehed rays. They rise between the paired dorsal

seutes and remain isolated from eaeh other, forming a series of dorsal finlets. In one of the photographed

speeimens, the last dorsal rays near the tail are not isolated but pressed together.

2 mm

SPIp.

Fig. 5. Gladiopycnodus karami gen. et sp. nov. The pelvie girdle as seen on the photo of a speeimen in

a private eolleetion.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

The anal fin begins with a very strong and extremely long spine that extends baekward greatly beyond

the tail. A few short, branehed rays follow this spine. The surfaee 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

dentieles. The spine is artieulated on the post-eoelomie bone and on the first pterygiophore, whieh is

longer and separated from the other anal pterygiophores.

5. The caudal skeleton (Fig. 2)

The eaudal skeleton is eovered by seales and thus remains unknown.

The small eaudal fin is not forked. Its posterior border is eonvex. The holotype has 17 prineipal rays, of

whieh the most external dorsal and ventral ones are segmented, pointed and a little shorter than the 15

others, that are segmented and branehed. There are 5 dorsal and 5 ventral proeurrent rays.

6. Squamation (Figs 5-6)

Gladiopycnodus karami gen. et sp. nov. possesses four different types of seutes and seales.

The dorsal margin of the fish is eovered by large and deep paired seutes that are ornamented with

tubereles and some small irregular ridges. The two last seutes are less deep than the more anterior pieees

of the series. The holotype bears thirteen pairs of these dorsal seutes. One sample seen on photo has

about twenty paired dorsal seutes. This dififerenee in the number of dorsal seutes perhaps refieets an

individual variation within the speeies, but eould have also a sexual origin. The ventral margin of the

eaudal region of the fish is devoid of sueh seutes.

The body is entirely eovered by small fiake-like seales that are irregularly imbrieated. Most seales have

a smooth posterior border, but some of them bear a few very small spines on their posterior border.

A series of large rounded seales eover the eaudal pedunele. They are ornamented with tubereles.

Two pelvie seutes and one posteloaeal seute are visible on the photo of the speeimen that shows the

pelvie girdle. A part of their margins is spiny.

SC c.

Fig. 6. Gladiopycnodus karami gen. et sp. nov., holotype CLC S-393. A. First dorsal seute.

B. Some seales of the body. C. One seale near the eaudal fin.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Genus Monocerichthys gen. nov.

um:lsid:zoobank.org:aet:2D642D4F-2B85-4838-90BD-B967E8345947

Type-species

Monocerichthys scheuchzei gen. et sp. nov. (by monotypy)

Diagnosis

As for the speeies (monospeeifie genus).

Etymology

From the Greek monokerds, unieom, and ichthys, fish.

Monocerichthys scheuchzeri gen. et sp. nov.

Figs 7-14

um:lsid:zoobank.org:aet:74512CQ9-6F58-4711-A028-93AFA9CBlQFF

Diagnosis

Gladiopyenodontid with the prefrontal elongated in a short rostrum, outpaeing only slightly the lower

jaw level. Prefrontal with a spiny anterior tip. Vomer bearing small rounded molariform teeth irregularly

ranged. Frontal short, narrow and with a weakly developed median protuberanee. Dermosupraoeeipital,

parietal and dermopterotie longer than deep and artieulated with a large supratemporal. Eong nuehal

horn with a broad basis, a very spiny posterior border and resting on the dermosupraoeeipital, the

supratemporal and the posttemporal. Orbitosphenoid artieulated with the mesethmoid. Well developed

triangular dermosphenotie. Premaxilla long, broad, toothless and sutured by its dorsal margin to the

prefrontal. Earge, toothless and triangular shaped maxilla. Small lower jaw. Hypertrophied trapezoid

preoperele eovering a great part of the eheek. Small operele with a rounded dorsal border and an

aeuminate ventral tip. Exposed part of the hyomandibula-dermohyomandibula eonsiderably smaller than

the preoperele. Very large peetoral girdle elosely assoeiated to the skull and forming a sort of eephalo-

thorax. Posttemporal artieulated with the supratemporal. Hypertrophied hypereleithrum. Strongly

hypertrophied eleithrum with a long and broad ventral braneh and a large posterior ventral proeess.

Eong peetoral spine artieulated on the eleithrum. 17 neural spines, all fused to their neural arehes, before

the epiehordal series. Neural and haemal spines short and broad. Posteoelomie bone obliquely oriented.

Short dorsal fin loeated in the middle of the baek, with 13 pterygiophores, 1 small spiny ray and 12

segmented rays. Very short anal fin. Caudal fin with a eonvex posterior margin and with 17 prineipal

rays, 6 dorsal and 7 ventral proeurrent rays. Body entirely eovered with very small flake-like seales. No

dorsal ridge seutes. One large posteloaeal seute below the posteoelomie bone.

Etymology

The speeies name of this new gladiopyenodontid fossil flsh is dedieated to Johann Jaeob Seheuehzer

(1672-1733), the Swiss physieian who was the first to illustrate a Eebanese fossil flsh in his hook Piscium

Queralae et Vindiciae (1708) [ef Gayet et al. 2012: 10].

Holotype and unique specimen

Sample CEC S-413a, b, the two faees of a eomplete speeimen (Figs 7-8). Total length: 102 mm.

Formation and locality

Marine Eate Cenomanian, Haqel, Eebanon.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

Holotype morphometric data

The morphometric data are given in % of the holotype standard length (87 mm).

Length of the head (opercle included).37.8 %

Length of the cephalo-thorax (cleithrum included). 66.7 %

Depth of the head (with the nuchal horn).37.3 %

Depth of the head (without the nuchal horn).64.7 %

Fig. 7. Monocerichthys scheuchzeri gen. et sp. nov. Holotype CLC S-413a (above) and b (below).

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Maximum depth of the body (just behind the nuehal horn).39.4 %

Eength of the peetoral spine. 19.9 %

Predorsal length.65.6 %

Basal length of the dorsal fin. 18.2 %

Preanal length.72.6 %

Basal length of the anal fin. 10.0 %

Depth of the eaudal pedunele.9.5 %

Osteology

1. The skull (Figs 9-10)

The dermal bones of the skull are eovered with small tubereles, but this ornamentation is not strongly

pronouneed.

The head is large, longer than high and triangular in shape. The snout is elongated in a short rostrum

formed by the long and broad prefrontal and by the long toothless premaxilla. The anterior tip of the

prefrontal is spiny. The rostrum outpaees only slightly the lower jaw level. The anterior region of the

long mesethmoid and the vomer are hidden by the prefrontal and the premaxilla. However, the most

anterior part of the premaxilla is broken on faee CEC S-413b and a part of the vomer is visible. The bone

bears very small, rounded molariform teeth, irregularly ranged.

The frontal is short, rather narrow and praetieally not eurved. It extends anteriorly beyond the orbit level

and exhibits a weakly developed median protuberanee loeated just before the orbit, but there is no frontal

horn. The posterior region of the skull roof is formed by the dermosupraoeeipital, and the paired parietals

and dermopteroties. These bones are rather small and longer than deep. There is no temporal fenestra.

A large supratemporal is sutured with the posterior margin of the dermosupraoeeipital, the parietal and

the dermopterotie. The autosphenotie is loeated below the dermopterotie. A long and strong pointed

Fig. 8. Monocerichthys scheuchzeri gen. et sp. nov. General reeonstruetion based on the two faees of

holotype CEC S-413a, b. Body seales are omitted.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

nuchal horn with a series of large spines on its posterior border is resting on the dermosupraoeeipital,

the supratemporal and also the posttemporal. The very broad basis of that horn extends on the beginning

of the baek.

The parasphenoid is very long and toothless. The orbitosphenoid is pressed against the mesethmoid. The

other endoehondral bones of the brainease are not aeeessible.

Fig. 9. Monocerichthys scheuchzeri gen. et sp. nov. Reeonstmetion of the skull and the peetoral girdle

based on the two faees of holotype CLC S-413a, b.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Fig. 10. Monocerichthys scheuchzeri gen. et sp. nov. The vomerian region of the skull in holotype CEC

S-413b.

Fig. 11. Monocerichthys scheuchzeri gen. et sp. nov. The dorsal fin of holotype CEC S-413a.

European Journal of Taxonomy 57: 1-30 (2013)

Small fragments of the metapterygoid and the entopterygoid are visible between the preoperele and the

parasphenoid, but a great part of those two bones remains hidden by the preoperele in the same way as

the quadrate and the sympleetie.

The upper jaw eontains a large, more or less triangular and toothless maxilla and a long, broad and

toothless premaxilla that is artieulated to the prefrontal by its upper margin. The lower jaw is triangular

in shape and rather small. The dentary is redueed to its ventral braneh. Its ventral margin is spiny. The

teeth are not preserved. Other gladiopyenodontid genera have two teeth on the dentary. There is a large

angular and a small artieular. It is not possible to determine whether the preartieular was toothed or not

or whether a marked eoronoid proeess was present. The preartieular bears molariform teeth on its inner

faee in some other gladiopyenodontid fishes.

The orbit is small. A large triangular dermosphenotie is loeated below the frontal and before the parietal

and the dermopterotie. No other orbital bone is preserved.

The exposed part of the hyomandibula-dermohyomandibula is important; however, it is mueh smaller

than the hypertrophied trapezoid-shaped preoperele, that almost entirely eovers the eheek. The operele

is a small bone, deeper than long, dorsally rounded and ventrally aeuminate. The suboperele and the

interoperele are absent, as in all pyenodontomorph fish.

The hyoid bar, the branehiostegal rays and the branehial skeleton are not visible.

2. The girdles (Figs 8-9)

The peetoral girdle is elosely assoeiated to the skull, both struetures forming a sort of eephalo-thorax.

The peetoral bones are ornamented with small tubereles. The posttemporal is a deep bone, narrow in its

upper part but ventrally broader, that is sutured anteriorly to the supratemporal and dorsally to the nuehal

horn. The hypereleithrum (= supraeleithrum) is hypertrophied in a large, more or less ovoid plate. The

Fig. 12. Monocerichthys scheuchzeri gen. et sp. nov. The anal fin of holotype CLC S-413a.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

eleithmm is gigantie, with a well developed dorsal braneh, a long and broad ventral braneh and a very

large posterior ventral proeess. The peetoral fin is replaeed by a long strong spine. There is no soft ray.

No pelvie bones are visible, but small fragments of rays from the ventral fins are preserved just above

the posterior point of the peetoral spine on faee CEC S-413a.

3. The axial skeleton (Fig. 8)

The trunk is fusiform and not deep-bodied as usual in most pyenodontomorph fishes. The abdominal

part of the axial skeleton progressively elevates and anteriorly reaehes the orbit level. The vertebrae are

eonstituted by the dorsal and ventral areoeentra only. The notoehord is not eompletely surrounded by

the arehes. There are 17 neural spines before the epiehordal series of the eaudal skeleton. They are short,

but rather broad. All those neural spines are fused to their neural arehes. Non of them are autogenous.

The first neural spine eorresponds to the first ossified neural areh. The fourth neural spine is fused to

two neural arehes. That is probably an individual variation and not a speeifie eharaeter. A few badly

preserved traees of the basiventrals from the abdominal region are visible, but the basiventrals of the

LEP

NEUR

NEUREP

Fig. 13. Monocerichthys scheuchzeri gen. et sp. nov. Reeonstruetion of the eaudal skeleton, based on the

two faees of holotype CEC S-413a, b.

European Journal of Taxonomy 57: 1-30 (2013)

caudal region and the haemal spines are lost, exeept the last ones near the tail. Four long and strong ribs

are seen just behind the hypereleithrum on faee CLC S-413b. The few preserved haemal spines are short

and broad, as the neural spines.

Only fragments of the most ventral part of the posteoelomie bone are preserved. The bone is well

developed, obliquely oriented and with an enlarged lower extremity that is direeted baekwards.

4. The dorsal and anal fins (Figs 8, 11-12)

The dorsal fin is short but rather high and is loeated in the middle of the baek. The fin begins with 1

small spiny ray, followed by 12 long segmented and branehed rays. There are 13 pterygiophores, eaeh

of them bearing one ray.

The anal fin is partly preserved on faee CLC S-413a but is eompletely missing on faee b. The fin is still

shorter than the dorsal one and is also loeated on the middle of the body, far from the tail. Fragments of

three pterygiophores and of four rays are visible in the beginning of the fin, just behind the posteoelomie

bone. Two very small pterygiophores and three very small rays are visible at the posterior extremity of

the fin.

5. The caudal skeleton (Fig. 13)

A eaudal pedunele is present. The eaudal skeleton is not very well preserved, but the two faees CLC

S-413a and b eomplete eaeh other and allow to entirely reeonstruet the whole strueture. The eaudal

fin is dorsally supported by 5 epiehordals and ventrally by 12 hypoehordals. The tenth and eleventh

hypoehordals are moderately broadened, but there is no real hypertrophy. The hypoehordal series

begins one vertebral segment before the epiehordal series. No urodermal is visible, but this is probably

Fig. 14. Monocerichthys scheuchzeri gen. et sp. nov. Body seales from holotype CLC S-413a, from the

dorsal region near the tail.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

a taphonomie artefaet. Indeed, a well developed urodermal does exist in some gladiopyenodontid fishes

more speeialized than M. scheuchzeri gen. et sp. nov. (pers. obs.).

The eaudal fin is not very large and has a eonvex posterior margin (Poyato-Ariza & Wenz 2002: fig. 36B).

There are 17 prineipal rays. The most dorsal and ventral prineipal rays are segmented but unbranehed.

The other prineipal rays are segmented and branehed. There are 6 dorsal and 7 ventral proeurrent rays.

6. Squamation (Fig. 14)

The body is entirely eovered with very small, flake-like seales that are imbrieated one into another.

Rather similar seales exist in Gladiopycnodus karami gen. et sp. nov., but they are deeper and mueh

larger. There are no peeuliar seales or seutes, either on the dorsal ridge or on the ventral keel.

A large irregular posteloaeal seute is present just below the posteoelomie bone.

Genus Rostropycnodus gen. nov.

um: lsid:zoobank.org: aet: 66711DD0-8E43 -4531-B64A-21071FE34C98

Type-species

Rostropycnodus gayeti gen. et sp. nov. (by monotypy)

Diagnosis

As for the speeies (monospeeifle genus).

Etymology

From the Eatin rostrum, rostrum. The generie name Pycnodus is added.

Rostropycnodus gayeti gen. et sp. nov.

Figs 15-21

um: Isid: zoobank, org: aet: 89707CF2-AF 8E-450E-8D6F -16A0B02EDE93

Diagnosis

Gladiopyenodontid with a very long snout, forming a rostmm anteriorly outpaeing the lower jaw level

(Fig. 18). Anterior tip of the rostmm formed by both the prefrontal and the premaxilla. Anterior extremity

of the prefrontal bearing four small spines. Frontal bearing one long pointed horn. Dermosupraoeeipital

forming a nuehal proeess. Small lower jaw. Preoperele mueh larger than the exposed part of the

hyomandibula-dermohyomandibula. Posttemporal artieulated with the nuehal proeess. Hypertrophied

eleithmm with a long and broad ventral braneh. Short and broad peetoral spine. Two large posteleithra.

Nuehal horn absent. 17 neural spines and 17 haemal spines before the epiehordal and hypoehordal

series. Dorsal fin with 6 pterygiophores. Anal fin with a long and strong initial spine and 4 short soft

rays. Caudal skeleton with 6 epiehordal and 7 hypoehordal elements. East three hypoehordal elements

moderately broadened. Caudal fin with 21 rays and a eonvex posterior margin. Body entirely eovered by

large, tubereulated seales with a spiny anterior border.

Etymology

The speeies name of the new gladiopyenodontid is dedieated to Mireille Gayet (Eyon), the well known

Freneh palaeontologist, who has greatly improved our knowledge of the Eate Cretaeeous iehthyofauna

of Eebanon.

European Journal of Taxonomy 57: 1-30 (2013)

Material examined

Holotype

Sample CLC S-608a, b, the two faees of a nearly eomplete speeimen (Fig. 15). The tip of the rostrum

and the eaudal fin are missing. Total length: 47.3 mm.

Fig. 15. Rostropycnodus gayeti gen. et sp. nov. Holotype CLC S-608a (above) and 608b (below).

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Paratypes

Sample CEC S-337, a eomplete speeimen (Fig. 16). The body and the tail are severely emshed. Total

length: 59.8 mm.

Sample CEC S-595, a nearly eomplete speeimen (Fig. 17). The tip of the rostrum is missing. Total

length: 58.6 mm.

Fig. 16. Rostropycnodus gayeti gen. et sp. nov. Paratype CEC S-337.

Fig. 17. Rostropycnodus gayeti gen. et sp. nov. Paratype CEC S-595.

European Journal of Taxonomy 57: 1-30 (2013)

Formation and locality

Marine Upper Cenomanian, Haqel, Lebanon.

Holotype morphometric data

Due to the ineompleteness of the available speeimens, no aeeurate morphologieal data ean be provided

and the reeonstmetion is therefore eomposite.

Osteology

1. The skull (Fig 19)

The head, rostrum and opereular series ineluded, is twiee as long as the body. The dermal bones of the

skull are ornamented with a few thin ridges and many tubereles.

The snout is elongated, forming a long rostrum that is, however, shorter than in Gladiopycnodus karami

gen. et sp. nov. The preorbital length, rostrum ineluded, represents 59.5 % of the total length of the

skull. The rostrum eonsists 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 gladiopyenodontid genera.

The frontal and the dermosupraoeeipital 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 whieh the anterior and posterior borders are spiny. The dermosupraoeeipital is enlarged and

forms a long and broad nuehal proeess. The autosphenotie, parietal and dermopterotie are rather small

bones. The parietal does not bear a brush-like proeess. The large supratemporal is pressed against the

dermosupraoeeipital.

The parasphenoid is long and toothless. A well developed orbitosphenoid is artieulated on the rear

of the mesethmoid. The other endoeranial bones of the brainease are hidden by the hyomandibula-

dermohyomandibula, the operele and the hypereleithrum.

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 loeated under the prefrontal and is sutured with

it all along its upper margin. The maxilla is small, eonsiderably shorter than the premaxilla, lanee head¬

shaped and posteriorly loeated.

The lower jaw is small, triangular in shape and, by far, does not reaeh the anterior extremity of the rostrum.

The short dentary bone, well visible on speeimen CLC S-608, is redueed to its ventral braneh. The teeth

are not preserved. The other bones of the mandible are too erushed to allow adequate deseription.

The bones of the palatoquadrate areh are hidden by the preoperele.

A large dermosphenotie is the only part preserved of the infraorbital series.

The exposed part of the hyomandibula-dermohyomandibula is mueh smaller than the large preoperele

that is, however, not as hypertrophied as in Gladiopycnodus karami gen. et sp. nov. The preoperele is

separated from the lower margin of the fish by the eleithrum, another differenee from G. karami gen.

et sp. nov.. The small operele is a deep ovoid bone with an aeuminate ventral extremity. The hyoid bar

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Fig. 18. Rostropycnodus gayeti gen. et sp. nov., paratype CEC S-337. Reeonstmetion based on the three

speeimens. Body seales omitted.

Fig. 19. Rostropycnodus gayeti gen. et sp. nov., paratype CEC S-337 Reeonstmetion of the skull based

on the three speeimens.

European Journal of Taxonomy 57: 1-30 (2013)

and the branchiostegal rays are not visible. A fragment of a branehial bone and a few short nail-shaped

branehiospines are preserved in the orbit of speeimen S-337.

2. The girdles (Figs 18-19)

The massive peetoral girdle is intimately assoeiated with the skull, forming a sort of eephalo-thorax.

The posttemporal is well developed and loeated just behind the hypereleithrum. The hypereleithrum

(= supraeleithrum) is a large triangular bone with some spines on its posterior border. The eleithrum

is really gigantie, with a very broad dorsal braneh and a very long, broad and pointed ventral braneh

reaehing the lower jaw level. There are two large posteleithra well visible on sample CLC S-337 and

loeated behind the eleithrum. The peetoral fin is replaeed by a short but very broad spine that is partially

fused with the eleithrum.

Two small pelvie bones are preserved between the eleithrum and the post-eoelomie bone on speeimen

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 eomplete vertebral eolumn. The notoehord is

ineompletely surrounded by the neural and haemal arehes, whieh are redueed. The neural and haemal

spines are well developed. There are 17 neural spines before the epiehordal series and 17 haemal spines

before the hypoehordal 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 inereases from

the fourth to sixth one and then deereases from the seventh to the seventeenth. The neural and haemal

spines are not interloeked together by interdigitating sutures. The presenee of ribs is uneertain, the situs

viscerum being hidden by the gigantie eleithrum.

The post-eoelomie bone is moderately long, robust, extremely broad and baekwardly eurved. Dorsally

it reaehes 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

eaeh other. The dorsal rays are lost but, eonsidering the spaeing 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

aeuminate posterior tip and a very broad basis artieulated on the post-eoelomie bone and the first anal

pterygiophore, whieh is mueh longer than the following ones. The pointed tip of the spine reaehes the

level of the tail but does not outpaee the eaudal 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. 21 A)

The eaudal skeleton is rather well preserved on holotype CLC S-608a, b. There are 6 epiehordal elements

and 7 hypoehordal elements. The last three hypoehordal pieees are moderately broadened. No urodermal

is visible.

Paratype CLC S-595 possesses a more or less eomplete eaudal fin, but essentially preserved as a print.

There are 21 rays, but it is not possible to distinguish the proeurrent from the prineipal rays. The fin has

a eonvex distal border (Poyato-Ariza & Wenz 2002: fig. 36 B).

6. Squamation (Fig. 2IB)

The body is entirely eovered by large deep seales that are imbrieated. Their entire surfaee is ornamented

with well developed tubereles and their anterior border is often irregularly spiny, as elearly seen on

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Fig. 20. Rostropycnodus gayeti gen. et sp. nov. The pelvie and anal region of holotype CEC S-608

(above) and of paratype CEC S-595 (below).

Fig. 21. Rostropycnodus gayeti gen. et sp. nov. A. Caudal skeleton of holotype CEC S-608a. The arrows

point on the more external proeurrent rays of the eaudal fin. B. Body seales of paratype CEC S-595.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

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 fianks. 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.

Discussion

1. Gladiopycnodus gen. nov., Monocerichthys gen. nov. and Rostropycnodus gen. nov. within

Neopterygii

Gladiopycnodus gen. nov., Monocerichthys gen. nov. din& Rostropycnodus gen. nov. exhibit some very

peculiar characters: (1) the mesethmoid is a long and massive bone including the lateral ethmoids, (2)

a dermosupraoccipital is present, (3) the preopercle is greatly hypertrophied and is sutured with the

hyomandibula, (4) the interopercle and subopercle are lost, (5) a dermohyomandibula is associated

with the hyomandibula, (6) the large prearticular bears molariform teeth, (7) the maxilla is plate-like

and toothless, (8) there is no supramaxilla, (9) the parasphenoid is elongated and toothless, (10) the

pelvic girdle is reduced, (11) a postcoelomic bone is present, (12) the neural and ventral arches are not

fused together and they do not entirely surround the notochord, and (13) there are dorsal, pelvic and

postcloacal scutes.

Within neopterygian fishes, only Pycnodontomorpha possess these thirteen characters associated. The

three new Lebanese genera can be confidently ranged in this superorder.

2. Gladiopycnodus gen. nov., Monocerichthys gen. nov. and Rostropycnodus gen. nov. within

Pycnodontomorpha

As already mentioned, Nursall (2010) divided Pycnodontomorpha [new usage for the former

Pycnodontiformes] into two orders, Gyrodontiformes and Pycnodontiformes [new usage for the former

Pycnodontoidei]. Gyrodontiformes are characterized by the presence of two dermosupraoccipitals, by a

dentary composed of two branches, one dorsal and one ventral, by their styliform teeth on the premaxilla

and the dentary, by their short parasphenoid and by their snout, cheek and gular region being completely

covered with small bony tesserae. In Pycnodontiformes, there is only one dermosupraoccipital,

the dentary has kept only its ventral branch, the dorsal one being lost, the teeth are incisiform, the

parasphenoid is long, and there are true infraorbitals.

The unique dermosupraoccipital, the dentary reduced to the ventral branch, the incisiform teeth

borne by the dentary, the long parasphenoid and the infraorbitals of Gladiopycnodus gen. nov.,

Monocerichthys gen. nov. and Rostropycnodus gen. nov. clearly show that these three fishes belong to

the Pycnodontiformes and not to the Gyrodontiformes.

3. Gladiopycnodus gen. nov., Monocerichthys gen. nov. and Rostropycnodus gen. nov. within

Pycnodontiformes

However, Gladiopycnodus gen. nov., Monocerichthys gen. nov. and Rostropycnodus gen. nov. differ

from all other known pycnodontiform fishes by their very peculiar snout anatomy and the presence

of a strong pectoral spine articulated with the cleithrum and replacing the lost pectoral fin. These two

apomorphies justify the erection of the new family Gladiopycnodontidae.

4. Gladiopycnodus gen. nov., Monocerichthys gen. nov. and Rostropycnodus gen. nov. within

Gladiopycnodontidae

Monocerichthys gen. nov. seems to be the less evolved gladiopycnodontid genus. The overall morphology

of its skull is still rather similar to that of a classic pycnodontiform fish. Other gladiopycnodontid genera

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

have a more speeialized skull, sometimes differing markedly from the usual pyenodontiform eranial

morphotype (pers. obs.). The rostrum of Monocerichthys gen. nov. is not very long and outpaees only

slightly the lower jaw level. This outpaeing is more pronouneed in the other members of the family

(pers. obs.).

On the other hand, Gladiopycnodus gen. nov. and Rostropycnodus gen. nov. belong to the more

speeialized members of Gladiopyenodontidae fam. nov. Indeed, these two Eebanese speeies had already

lost the nuehal horn present in the primitive genera of the family (pers. obs.) and they share three

apomorphies not present in primitive members of the lineage, i.e., an extremely elongated rostrum, a

greatly hypertrophied anal spine and the first anal pterygiophore being mueh longer than the following

ones.

5. Gladiopycnodontidae fam. nov. within Pycnodontiformes

We follow hereafter the phytogeny ofPyenodontiformes established on eranial and posteranial eharaeters

by Poyato-Ariza & Wenz (2002: fig. 43; 2004: fig. 15; 2005: fig. 10). The phytogeny presented by Nursall

(1996b: fig. 4, 18) is less detailed and less eomplete. Kriwet (2005: 62) also proposed a phytogeny of

these fishes differing from that of Poyato-Ariza & Wenz (2002), but being based only on eranial data.

Brembodidae are eonsidered as the most primitive lineage within Pyenodontiformes. Data on this family

eome from Tintori (1980), Nursall (1999) and Poyato-Ariza & Wenz (2002). Brembodidae have a deep

body, with a dorsal gibbosity or a dorsal spiny expansion formed by the nuehal and dorsal seutes. As

in Gyrodontiformes, they have bony tesserae in the gular region and their eheek is eompletely eovered,

but here by a greatly hypertrophied first infraorbital and not by bony tesserae. Their premaxilla bears

3 teeth and their dentary 4 or 5 teeth. Their unpaired fins are bordered by fringing fiilera. Their body is

eompletely eovered by seales as in Gyrodontiformes.

Macromesodon Blake, 1905 is another primitive pyenodontiform fish with a dorsal angle-shape

prominenee. Woodward (1918) and Poyato-Ariza & Wenz (2004) provide information on the

genus. Macromesodon has tubular infraorbitals, but still possesses bony tesserae on the eheek, as in

Gyrodontiformes, and 4 teeth on the dentary. The tesserae of the gular region are lost. The body is

eovered by seales only in its anterior region and not in its eaudal part. The fringing fiilera on the unpaired

fins are lost. These three last eharaeters show that Macromesodon oeeupies a more apomorphie position

in the pyenodontiform phylogenetie tree than Brembodidae.

Gladiopyenodontidae fam. nov. share with more evolved Pyenodontiformes at least three eharaeters

that plaee them in a more advaneed position in the phytogeny of the order than Brembodidae and

Macromesodon. The infraorbitals are redueed and eover only a small part of the eheek, whieh is devoid

of bony tesserae. The number of teeth on the dentary is redueed (generally 2). There are only two

branehiostegal rays that are separated from eaeh other (pers. obs.).

Pyenodontidae are the largest and most advaneed family within Pyenodontiformes. They eontain

about twenty genera. The most primitive of them is Akromystax Poyato-Ariza & Wenz, 2005 from the

Cenomanian of Eebanon (Poyato-Ariza & Wenz 2005: fig. 10). The most speeialized members of the

family are grouped in the subfamily Nursalliinae (Poyato-Ariza & Wenz 2002: fig. 43). The prineipal

apomorphy of Pyenodontidae is the presenee of a brush-like proeess on the parietal, a strueture absent

in less evolved Pyenodontiformes and in Gladiopyenodontidae fam. nov.

The systematie position of Gladiopyenodontidae fam. nov. is thus between Brembodidae and

Macromesodon on the one hand, and Pyenodontidae on the other hand, a region of the pyenodontiform

European Journal of Taxonomy 57 : 1-30 ( 2013 )

phylogenetic tree containing only the family Coccodontidae (including Trewavasiidae), another highly

specialized pycnodontiform lineage from Lebanon.

6. Gladiopycnodontidae fam. nov., Coccodontidae and Coccodontoidea superfam. nov.

A comparison between Gladiopycnodontidae fam. nov. and Coccodontidae is thus necessary to define

the possible relationships between the two families.

As already said, Monocerichthys gen. nov. is the least specialized gladiopycnodontid genus. On the other

hand, Coccodus Pictet, 1850 is the least specialized member of the family Coccodontidae (Capasso et

al. 2010: 160-161). Data on Coccodus hereafter mentioned come from Pictet (1850), Davis (1887),

Hay (1903), Poyato-Ariza & Wenz (2002), Kriwet (2005) and principally from our own observations

(Taveme & Capasso, in prep.).

Monocerichthys gen. nov. and Coccodus share a series of characters not present in the other pycnodontiform

families: (1) the body is fusiform and without dorsal or ventral apex, (2) the frontal exhibits a small

median protuberance, (3) the large supratemporal is articulated with the dermosupraoccipital, the parietal

and the dermopterotic, (4) the large posttemporal is sutured to the supratemporal, (5) the very large

pectoral girdle is closely associated to the skull, forming a sort of cephalo-thorax, (6) the hypercleithrum

is hypertrophied, (7) the cleithrum is greatly hypertrophied with a long and broad ventral branch and a

large ventral posterior process, (8) there are less than 20 vertebral segments before the epichordal series,

(9) the neural and haemal spines are short, (10) the dorsal fin is short and located in the middle of the

back, (11) the anal fin is short and largely separated from the tail and (12) there is never more than one

postcloacal scute.

These twelve characters thus can be used to define the new super-family Coccodontoidea superfam.

nov., grouping Gladiopycnodontidae fam. nov. and Coccodontidae.

However, in spite of all these resemblances, Coccodontidae and Gladiopycnodontidae fam. nov. cannot

be confounded. They greatly differ at the level of their snout morphology, that of Coccodontidae

remaining typically pycnodontid. Coccodontidae have also retained a normal pectoral fin with soft rays,

whereas Gladiopycnodontidae fam. nov. have the pectoral fin replaced by a strong spine.

Acknowledgements

We thank Dr. Andrea Pessina, Superintendant of the Soprintendenza per i Beni Archeologici delT

Abruzzo - Chieti, for allowing us to study the fossil fishes of Luigi Capasso’s collection. We are grateful

to M. Luciano Lullo of the University of Chieti and to M. Hugo De Potter of the Royal Institute of

Natural Sciences of Belgium for their technical help. We also thank the anonymous colleagues who have

accepted to review our manuscript.

References

Capasso L.L., Abi Saad P. & Taveme L. 2009. Nursallia tethysensis sp. nov., a new pycnodont fish

(Neopterygii: fHalecostomi) from the Cenomanian of Lebanon. Bulletin de llnstitut Royal des Sciences

Naturelles de Belgique, Sciences de la Terre 79: 117-136.

Capasso L.L., Taveme L. & Nohra A.R 2010. A re-description of Hensodon spinosus, a remarkable

coccodontid fish (Actinopterygii, tPycnodontiformes) from the Cenomanian (Late Cretaceous) of

Haqel, Lebanon. Bulletin de I Tnstitut Royal des Sciences Naturelles de Belgique, Sciences de la Terre

80: 145-162.

TAVERNE E. & CAPASSO E., Gladiopycnodontidae fam. nov. (Pisces) from the Cretaeeous of Eebanon

Davis J.W. 1887. The fossil fishes of the Chalk of Mount Eebanon, in Syria. Scientific Transactions of

the Royal Dublin Society series 2, 3(12): 457-636.

Davis J.W. 1890. On a new speeies of Coccodus (C. lindstroemi, Davis). The Quaterly Journal of the

Geological Society of London 46: 565-568.

Forey P.E., Eu Y., Patterson C. & Davies C.E. 2003. Fossil fishes of the Cenomanian (Upper Cretaeeous)

of Namoura, Eebanon. Journal of Systematic Palaeontology 1(4): 227-330. http:// dx. doi.org/10.1017/

S147720190300107X

Gayet M. 1984. Ichthyoceros spinosus nov. gen., nov. sp., du Cenomanien inferieur de Hakel (Eiban) et

ses affinites avee le geme Trewavasia (Pisees, Pyenodontiformes, Coeeodontidae). Bulletin du Museum

National d’Histoire Naturelle 4® serie, 6, seetion 6, 3: 287-307.

Gayet M., Abi Saad P. & Gaudant O. 2012. Les fossiles du Liban. Memoire du temps. Ed. Desiris, Gap.

Hay O.P 1903. On a eolleetion of Upper Cretaeeous fishes from Mount Eebanon, Syria, with deseriptions

of four new genera and nineteen new speeies. Bulletin of the American Museum of Natural History

19(10): 395-452.

Kriwet J. 2001. Palaeobiogeography of pyenodontiform fishes (Aetinopterygii, Neopterygii). In\

Melendez G., Herrera Z., Delvene G. & Azanza B. (eds) Los fdsilesy lapaleogeographia. XII Jornadas

de la Sociedad Espahola de Paleontologia: 121-130. Universidad de Zaragoza, Zaragoza.

Kriwet J. 2004. A new pyenodont fish genus (Neopterygii: Pyenodontiformes) from the Cenomanian

(Upper Cretaeeous) of Mount Eebanon. Journal of Vertebrate Paleontology 24(3): 525-532. http://

dx.doi.org/l0.l67l/0272-4634r2004l024[0525:ANPFGN12.0.CO:2

Kriwet J. 2005. A eomprehensive study of the skull and dentition of pyenodont fishes. Zittelliana A45:

135-188.

Nursall J.R. 1996a. Distribution and eeology of pyenodont fishes. In\ Arratia G. & Viohl G. (eds)

Mesozoic Fishes - Systematics and Paleoecology: 115-124. Verlag Dr. F. Pfeil, Mtinehen.

Nursall J.R. 1996b. The phylogeny of pyenodont fishes. In\ Arratia G. & Viohl G. (eds) Mesozoic Fishes

-Systematics and Paleoecology. 125-152. Verlag Dr. F. Pfeil, Mtinehen.

Nursall J.R. 1999. The family fMesturidae and skull of the pyenodont fishes. In\ Arratia G. & Viohl G.

(eds) Mesozoic Fishes 2 - Systematics and Paleoecology. 153-188. Verlag Dr. F. Pfeil, Mtinehen.

Nursall J.R. 2010. The ease for pyenodont fishes as the fossil sister-group of teleosts. In\ Nelson J.S.,

Sehultze H.-P. & Wilson M.V.H. (eds) Origin and Phylogenetic Interrelationships of Teleosts'. 37-60.

Verlag Dr. F. Pfeil, Mtinehen.

Nursall J.R. & Capasso E. 2004. Gebrayelichthys (novum), an extraordinary genus of neopterygian fishes

from the Cenomanian of Eebanon. In'. Arratia G. & Tintori A. (eds) Mesozoic Fishes 3 - Systematics,

Paleoenvironments and Biodiversity. 317-340. Verlag Dr. F. Pfeil, Mtinehen.

Nursall J.R. & Capasso E. 2008. Additional speeimens from Eebanon reveal more of the strueture of the

pyenodont fish Trewavasia carinata (Davis, 1887). In'. Arratia G., Sehultze H.-P. & Wilson M.V.H. (eds)

Mesozoic Fishes 4 - Homology and Phylogeny. 143-166. Verlag Dr. F. Pfeil, Mtinehen.

Pietet F.-J. 1850. Description de quelques poissons fossiles du Mont Liban. Imprimerie J.-G. Fiek,

Geneve.

Poyato-Ariza F.J. & Wenz S. 2002. Anew insight into pyenodontiform fishes. Geodiversitas 24(1): 139-

248.

European Journal of Taxonomy 57 : 1-30 ( 2013 )

Poyato-Ariza FJ. & Wenz S. 2004. The new pyenodontid fish genus Turbomesodon, and a revision of

Macromesodon based on new material from the Lower Cretaeeous of Las Hoyas, Cuenea, Spain. In\

Arratia G. & Tintori A. (eds) Mesozoic Fishes 3 - Systematics, Paleoenvironments and Biodiversity.

341-378. Verlag Dr. F. Pfeil, Munehen.

Poyato-Ariza F. J. & Wenz S. 2005. Akromystax tilmachiton gen. et sp. nov., a new pyenodontid fish from

the Lebanese Late Cretaeeous of Haqel and En Nammoura. Journal of Vertebrate Paleontology 25(1):

27-45. http://dx.doi.org/10.1671/0272-4634t20051025r0027:ATGESN12.0.CO:2

Tintori A. 1980. Two new pyenodonts (Pisees, Aetinopterygii) from the Upper Triassie of Eombardy (N.

Italy). Rivista Italiana di Paleontologia e Stratigrafia 86(4): 795-824.

Woodward A.S. 1918. The Fossil Fishes of the English Wealden and Purbeck Formations. Part 2\ 49-

104. Palaeontologieal Soeiety, Eondon.

Manuscript received: 19 March 2013

Manuscript accepted: 5 August 2013

Published on: 2 October 2013

Topic editor: Christian de Muizon

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium; Natural History

Museum, Eondon, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.