European Journal of Taxonomy 101: 1-27

http://dx.doi.org/10.5852/ejt.2014.101

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2014 • Taveme L. & Capasso L.

This work is licensed under a Creative Commons Attribution 3,0 License.

Research article

um:lsid:zoobank.org:pub:C320F2F8-E9BB-46FC-9A0C-56ECF4BlCE2B

On the “ Coccodus ” lindstroemi species complex (Pycnodontiformes,

Gladiopycnodontidae) from the marine Late Cretaceous of Lebanon,

with the description of two new genera

Louis TAVERNE 1 & Luigi CAPASSO 2

1 Royal Belgian Institute of Natural Sciences of Belgium, Directorate Earth and History of Life,

Rue Vautier 29, B-1000 Brussels, Belgium. E-mail: [email protected]

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

1-661000 Chieti, Italy. E-mail: [email protected]

1 urn:lsid:zoobank. org: author: OCF 81641 - 1DD1-4CBD-973 5-F1FE7EB0BCF 5

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

Abstract. The osteology of “Coccodus” lindstroemi is studied in detail and it is demonstrated that this

species does not belong to the genus Coccodus , but is a rather primitive member of the pycnodontiform

family Gladiopycnodontidae. Indeed, the snout of “ Coccodus ” lindstroemi is elongated in a rostrum

formed by the prefrontal and the premaxilla. This rostmm extends beyond the lower jaw level. The

toothless premaxilla is sutured by its upper margin to the lower margin of the long and broad prefrontal.

The pectoral fin is lost and replaced by a pectoral spine which articulates on the cleithrum. A long nuchal

spine resting on the dermosupraoccipital is present. The body is entirely covered by scales that are flake¬

like in the abdominal region and scute-like in the caudal region. Joinvillichthys gen. nov. is thus erected

with “ Coccodus ” lindstroemi as the type species. It is also shown that specimens with dumpier head and

body, usually ranged in “ Coccodus ” lindstroemi , represent another species of the same genus for which

the taxon Joinvillichthys kriweti gen. et sp. nov. is created. Specimens sometimes considered as possible

juveniles of “ Coccodus ” lindstroemi form a distinctive new genus and species of gladiopycnodontid

fish, Pankowskichthys libanicus gen. et sp. nov. Pankowskichthys differs from Joinvillichthys by many

osteological structures.

Key words. Pycnodontiformes, osteology, relationships, marine Cenomanian, Lebanon

Taverne L. & Capasso L. 2014. On the “ Coccodus ” lindstroemi species complex (Pycnodontiformes,

Gladiopycnodontidae) from the marine Late Cretaceous of Lebanon, with the description of two new genera.

European Journal of Taxonomy 101: 1-27. http://dx.doi.org/10.5852/ejt.2014.101

Introduction

The recently erected family Gladiopycnodontidae is a highly specialized lineage of the fossil fish order

Pycnodontiformes (Taverne & Capasso 2013). The family is endemic to the marine Late Cretaceous

of Lebanon and contains about ten genera (pers. obs.) most of them still undescribed. They are small

fishes with a large head and a rather fusiform body. Their snout ends in a rostrum that is formed by the

European Journal of Taxonomy 101: 1-27 (2014)

enlarged prefrontals and premaxillae that are sutured together. The pectoral fin is lost and replaced by

a strong pectoral spine. Some genera bear frontal, occipital and/or nuchal horns. The pectoral girdle

is hypertrophied and closely associated to the skull, forming a kind of cephalo-thorax. Three genera

belonging to this family have previously been described: Gladiopycnodus Taverne & Capasso, 2013,

Monocerichthys Taverne & Capasso, 2013 and Rostropycnodus Taverne & Capasso, 2013.

“ Coccodus ” lindstroemi Davis, 1890 is the first member of this family ever described. The original

description was based on a single complete specimen from Haqel, Tebanon, which has an elongated

head, a pointed snout and a long nuchal horn (Davis 1890: pi. 22). The fish was erroneously (Poyato-

Ariza & Wenz 2002: 145) considered as a new species of the already known Tebanese genus Coccodus

Pictet, 1850. Tater, other specimens resembling the holotype but with a dumpier head and body were

also referred to the species “C.” lindstroemi (Hay 1903: pi. 29, fig. 1; Gayet et al. 2012: 87, upper left

photo). More recently, small samples with an extremely long nuchal horn were considered as possible

juveniles of this species (Gayet et al. 2012: 87, upper right photo).

It should be noted that both the original description of “C.” lindstroemi by Davis (1890) and the

redescription by Hay (1903) give practically no informative data on the skull and the caudal skeleton of

the fish. Today, the osteology and the relationships of “C.” lindstroemi remain almost unknown. Poyato-

Ariza & Wenz (2002: 145) considered this fossil fish as “in need of revision” and doubted its taxonomic

assignment to the Pycnodontiformes.

The aims of the present paper are, firstly, to provide for the first time an adequate osteological description

of the concerned fishes, secondly, to demonstrate that they are true Pycnodontiformes and members of

the family Gladiopycnodontidae, and, thirdly, to show that three different species coexist within the

“C.” lindstroemi complex as understood today by some authors, the true “C.” linsdtroemi , the dumpier

species and the species comprising the so-called juveniles. The first two species are allocated to a first

new genus, the third species is placed in a second new genus.

Material and methods

The material belongs to the collections of the Royal Belgian Institute for Natural Sciences (IRSNB), the

Swedish Museum of Natural History (NRM), the American Museum of Natural History (AMNH) and

the Capasso-registered collection (CLC) in Chieti.

The specimens have been studied with Wild M 5 and Teica Wild M 8 stereo-microscopes. The figures

were drawn by the first author (L.T.) with a camera lucida. Aspersions with ethanol were used to improve

some observations.

List of abbreviations used in text-figures

= angular

ART

= articular

ASPH

= autosphenotic

= basioccipital

BRSTG

= branchiostegal rays

BSPH

= basisphenoid

CHY

= ceratohyal

CTT

= cleithrum

DHYOM

= dermohyomandibula

= dentary

DPTE

= dermopterotic

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

DSOC

dermosupraoccipital

DSPH

dermosphenotic

ENPT

entopterygoid (= endopterygoid)

EPCO 1-6

epichordals 1 to 6

frontal

HAEM

haemal arch

HAEMEP

haemal spine

HCLT

hypercleithrum (= supracleithrum)

HYCO 1-8

hypochordals 1 to 8

HYOM

hyomandibula

HYP

hypural plate (= fused hypochordals)

IORB 1

infraorbital 1

LEP

lepidotrichium (= ray)

METH

mesethmoid

MPT

metapterygoid

maxilla

NEUR

neural arch

NEUREP

neural spine

nuchal horn

opercle

OSPH

orbitosphenoid

parietal

PCLT

postcleithrum

PCOEL

postcoelomic bone

PELV

pelvic bone

PMX

premaxilla

POP

preopercle

PRART

prearticular

PRFR

prefrontal (= lateral dermethmoid ?)

PRO

prootic

parasphenoid

PSPH

pleurosphenoid

posttemporal

quadrate

RAD

pterygiophores (= radials)

ribs

SC b.

body scales

SCL

sclerotic bone

SCUd.

scutes of the dorsal ridge

SCUv.

scutes of the ventral keel

SPI

pectoral spine

supratemporal

symplectic

urodermal

vomer

f. V

foramen of the trigeminal nerve (V)

left

right

European Journal of Taxonomy 101: 1-27 (2014)

Results

Systematic paleontology

Subclass Actinopterygii Klein, 1885

Series Neopterygii Regan, 1923

Division Halecostomi Regan, 1923 sensu Patterson 1973

Superorder Pycnodontomorpha Nursall, 2010

Order Pycnodontiformes Berg, 1937 sensu Nursall 2010

Superfamily Coccodontoidea Taverne & Capasso, 2013

Family Gladiopycnodontidae Taverne & Capasso, 2013

Joinvillichthys gen. nov.

um:lsid:zoobank.org:act:6D820D8E-DF5C-41E9-B351-F46F7A9D4119

Type species: Coccodus lindstroemi Davis, 1890 (here designated).

Diagnosis

Gladiopycnodontid with an elongate prefrontal forming a short rostrum outpacing the lower jaw level.

Anterior extremity of the prefrontal acuminate and spiny. Vomer bearing small rounded molariform

teeth irregularly ranged. Orbitosphenoid, pleurosphenoid and basisphenoid present. Supratemporal

sutured to the rear of the skull. Premaxilla long, broad, toothless and sutured by its upper margin to the

prefrontal. Dentary bearing 2 incisiform teeth. Hypertrophied trapezoid preopercle covering the cheek.

Large first infraorbital (only known in Joinvillichthys kriweti gen. et sp. nov.). Exposed part of the

hyomandibula-dermohyomandibula much smaller than the preopercle. Long nuchal horn with a spiny

posterior margin and articulated only on the dennosupraoccipital. Pectoral girdle closely associated to

the skull, forming a cephalo-thorax. Cleithrum hypertrophied, with a gigantic posterior ventral process.

Hypercleithrum hypertrophied. Well developed posttemporal, with an acuminate posterior extremity. 16

to 17 neural spines, all fused to the neural arches, before the epichordal series. 10 haemal spines before

the hypochordal series (only known in Joinvillichthys lindstroemi). Short dorsal fin with 8 or 9 rays.

Short anal fin with 7 to 9 rays. 7 to 9 spiny scutes in the dorsal ridge. 3 or 4 scutes in the ventral keel, the

last scute or the two last ones associated to the postcoelomic bone. Body completely covered by small,

flake-like scales in the abdominal region and by large, scute-like scales in the caudal region.

Etymology

The generic name is chosen in memory of Lord Jean de Joinville (1224-1317), seneschal of Champagne,

who related in his biography of Louis IX the presentation of some Lebanese fossil fishes to this French

king at Saida during the seventh crusade (Gayet et al. 2012: 8). The Greek word ichthys, fish, is added.

Joinvillichthys lindstroemi (Davis, 1890)

Figs 1-11

Diagnosis

Joinvillichthys with a body depth comprising between 23.8 and 34.0 % of the standard length. No dorsal

prominence on the frontal. Maxilla triangular in shape. Small parietal. Dermosupraoccipital sutured with

the parietal and the dermopterotic. Dermopterotic deeper than long. Large dermosphenotic. Thin, rod¬

like opercle. Anterior ventral branch of the cleithrum present. T hin pectoral spine articulated on the rear

of the cleithral posterior process. Caudal fin with a convex posterior margin.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

Synonymy

Coccodus Lindstroemi Davis, 1890: 567, pi. 22.

“ Coccodus ” lindstroemi - Poyato-Ariza & Wenz, 2002: 145.

Holotype

LEBANON: sample NRM PZ P. 2073, a complete specimen from Haqel (Fig. 1), total length: 76 mm.

Other specimens

LEBANON: sample IRSNB N° P 9276, a nearly complete specimen (the caudal fin is missing) from

Hgula (Fig. 2), length: 93 mm; sample CLC S-138, a nearly complete specimen (a part of the caudal fin

is missing) from Haqel (Fig. 3), length: 82 mm; sample CLC S-324, a complete specimen from Haqel,

total length: 76 mm

Formation and locality

Marine Upper Cenomanian, Haqel and Hgula, Lebanon.

Morphometric data (Fig. 4)

The morphometric data are given in % of the standard length for the holotype NRM PZ P. 2073 (64 mm )

and for sample IRSNB N° P 9276 (93 mm). These two specimens represent the two extremes of the

species morphometric variation, as measured on the four studied samples

Length of the head (dermosupraoccipital included)

Holotype

.. 55.8 %

P9276

56.3 %

Fig. 1 . Joinvillichthys lindstroemi (Davis, 1890). Holotype, NRM PZ P. 2073.

European Journal of Taxonomy 101: 1-27 (2014)

Fig. 2. Joinvillichthys lindstroemi (Davis, 1890). Sample IRSNB N° P 9276.

Fig. 3. Joinvillichthys lindstroemi (Davis, 1890). Sample CLC S-138.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

Fig. 4. Joinvillichthys lindstroemi (Davis, 1890). General reconstruction based on holotype, NRM PZ

P 2073 and samples IRSNB N° P 9276, CLC S-138 and CLC S-324. The scale refers to sample CLC

S-138.

Fig. 5. Joinvillichthys lindstroemi (Davis, 1890). Skull and pectoral girdle of holotype, NRM PZ P.

2073.

European Journal of Taxonomy 101: 1-27 (2014)

Length of the cephalo-thorax (cleithrum included).63.9 % 68.1 %

Depth of the head (without the nuchal horn).35.4 % 25.0 %

Length of the nuchal horn.25.2 % 25.6 %

Maximum depth of the body (just behind the nuchal horn).34.0 % 23.8 %

Predorsal length.74.8 % 73.1 %

Basal length of the dorsal fin. 9.5 % 10.6 %

Preanal length.72.1% 73.7 %

Basal length of the anal fin. 8.2 % 9.4 %

Depth of the caudal peduncle. 5.4 % 5.3 %

The important individual differences in the values of the head and body depths are probably due to

sexual or seasonal variations.

Osteology

1. The skull (Figs 5-8)

The head is very large. Its length, from the tip of the snout to the basis of the nuchal horn, is equal to the

body length. According to the specimens, its depth, from the upper margin of the dermosupraoccipital

to the lower margin of the cleithrum, represents from 48 to 67 % of its length. The dermal bones of the

skull are ornamented with small tubercles.

Fig. 6. Joinvillichthys lindstroemi (Davis, 1890). Reconstruction of the skull and the pectoral girdle

based on holotype, NRM PZ P. 2073, and samples IRSNB N° P 9276, CLC S-138 and CLC S-324. The

scale refers to sample CLC S-138.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

The long, pointed rostrum slightly outpaces the lower jaw and is formed by two large paired bones, the

prefrontal and the premaxilla. The anterior tip of the prefrontal bears two or three very small spines.

Posteriorly, the bone reaches the orbit level. Only the most posterior part of the mesethmoid is visible.

The vomer is completely hidden by the premaxilla. However, a small anterior fragment of the premaxilla

is lost on sample CLC S-138 and a part of the vomer is visible. The bone bears small, rounded molariform

teeth that are irregularly ranged.

The frontal is short, not curved and slightly broader posteriorly than anteriorly. The posterior portion of

the frontal outpaces the level of the orbit. The posterior lateral part of the skull roof is formed on each

side by four small bones, the parietal, the dermosphenotic, the dermopterotic and the supratemporal.

The dermosphenotic partly covers the autosphenotic. The dermopterotic is deeper than long. The

supratemporal is sutured to the dermopterotic and does not reach the parietal. The dermosupraoccipital

occupies the median posterior part of the skull roof. This large bone is sutured with the frontal, the parietal,

the dermopterotic and the supratemporal. Along pointed nuchal horn is fixed to the dermosupraoccipital.

This horn is ornamented with long and thin striations and bears a series of spines on its posterior border.

Sample IRSNB N° P 9276 clearly shows the orbitosphenoid, the pleurosphenoid and the basisphenoid

in the orbit. The three bones are pressed against the frontal. The orbitosphenoid reaches anteriorly the

mesethmoid. The small basisphenoid is divided in a dorsal meningost and a short ventral belophragm.

Fig. 7. Joinvillichthys lindstroemi (Davis, 1890). Vomerian region of sample CLC S-138.

European Journal of Taxonomy 101: 1-27 (2014)

The parasphenoid is very long, almost straight, but it does not reach the posterior border of the skull that

is occupied by the basioccipital, as seen on the same specimen. Sample P 9276 also shows the very small

prootic with a large foramen for the trigeminal nerve (V) in its anterior border.

The anterior margin of the metapterygoid and the entopterygoid is visible between the preopercle and

the parasphenoid. The quadrate and the symplectic remain hidden by the preopercle and the cleithrum.

The premaxilla and the maxilla compose the upper jaw. As in other pycnodontomorph fishes, there

is no supramaxilla. The broad, long and toothless premaxilla is located below the prefrontal to which

it is sutured by its upper margin. The maxilla is large, toothless and triangle-shaped. The lower jaw

comprises the dentary, the prearticular, the angular and the articular. The articulation with the quadrate

is located at the level of the anterior border of the orbit. The prearticular is the largest bone of the series

but is partly covered by the maxilla and the preopercle. The teeth of the prearticular are not visible. The

articular and the angular are small bones. The dentary bears two incisiform teeth and is reduced to its

ventral branch. Its lower margin is denticulated.

The orbit is large and longer than deep. No orbital bone is preserved, except the dermosphenotic that is

sutured to the frontal, the parietal and the dermopterotic. Fragments of a sclerotic bony ring are visible

on the holotype.

The hyomandibula-dermohyomandibula and the preopercle are sutured together. The exposed part of the

hyomandibula-dermohyomandibula is much smaller than the greatly enlarged preopercle. The opercle is

a long and very thin bone wedged between the preopercle and the cleithrum.

Small fragments of branchial bones with a few long and acuminate branchiospines are visible on sample

IRSNB N° P 9276.

PRFR

METH

2 mm

PSPH

BSPH

OSPH

Fig. 8. Joinvillichthys lindstroemi (Davis, 1890). Orbital region of sample IRSNB N° P 9276.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

Fig. 9. Joinvillichthys lindstroemi (Davis, 1890). Caudal skeleton of sample IRSNB N° P 9276.

HYCO 8?

Fig. 10. Joinvillichthys lindstroemi (Davis, 1890). Hypochordal elements of sample CLC S-138. The

arrows indicate the position of the most external dorsal and ventral procurrent ray of the caudal fin.

European Journal of Taxonomy 101: 1-27 (2014)

2. The girdles (Figs 4-6)

As in all Gladiopycnodontidae, the enlarged pectoral girdle is closely associated to the skull, forming a

sort of cephalo-thorax. The dermal bones are ornamented with small tubercles. The postemporal rests

on the large ovoid hypercleithrum (= supracleithrum) and is articulated with the dermosupraoccipital

by its broad anterior border. Its posterior extremity is acuminate. The cleithrum is enormous, with a

well developed anterior branch and a very wide posterior process. There is a small postcleithrum. The

pectoral fin is replaced by a long and thin spine that is decorated with a few ridges and tubercles. The

spine is articulated on the rear of the cleithral posterior process.

No trace of a pelvic girdle is visible. It is possible that reduced pelvic bones and fins were present but

hidden by the gigantic cleithral posterior process. Such a situation exists in other gladiopycnodontid

fishes (Taverne & Capasso 2013: figs 8, 18).

2 mm

SCUd.

SCU d.

SCUv.

Fig. 11. Joinvillichthys lindstroemi (Davis, 1890). A. Dorsal ridge scutes of sample IRSNB N° P 9276.

B. Anterior flake-like body scales of sample CTC S-138. C. Posterior scute-like body scale of sample

CLC S-138. D. Ventral keel scutes of sample CLC S-138.

TAVERNE L. & CAPASSO F., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

3. The axial skeleton (Fig. 4)

As in most Gladiopycnodontidae, the trunk is fusiform and not deep-bodied. Sample IRSNB N° P 9276

has lost the scales on the body and so the well preserved axial skeleton is completely accessible. The

vertebrae are constituted by the dorsal and ventral arcocentra. They surround almost completely the

notochord. There are 17 neural spines before the epichordal elements and 10 haemal spines before the

hypochordal series. Before the level of the dorsal fin, the neural spines are long and narrow. Posteriorly,

the neural spines are much shorter but also a little broader. The haemal spines are short and broad. The

number of ribs is not determinable but ribs are present under the cleithral posterior process, as seen on

sample CLC S-138. The last ribs are very short. The postcoelomic bone is backwardly oriented and is

articulated with the ventral arcocentrum preceding the one bearing the first haemal spine. The bone is

broader ventrally than dorsally.

4. The dorsal and anal fins (Fig. 4)

The dorsal and anal fins are short and located in the middle of the body length. There are 8 or 9 dorsal

pterygiophores and also 8 or 9 anal pterygiophores, each of them bearing a ray. The first dorsal and anal

ray is spiny. The other rays are segmented.

5. The caudal skeleton (Figs 9-10)

Sample IRSNB N° P 9276 presents the best preserved caudal skeleton. The caudal peduncle is long and

includes 5 or 6 vertebral segments. There are 6 epichordals and a least 8 hypochordals. The hypochordals

are broader than the long, thin and pointed epichordals. In specimen IRSNB N° P 9276, the sixth and

seventh hypochordals are moderately broadened and partly fused together. In sample CFC S-138, the

sixth and seventh hypochordals are not fused and the broadening only exists on the seventh element.

The caudal fin has a convex posterior margin (Poyato-Ariza & Wenz 2002: fig. 36B) and contains 17 or

18 principal caudal rays. There are a few procurrent rays in each lobe.

6. Squamation (Fig. 11)

The body is entirely covered by scales ornamented with tubercles and imbricated one into another.

Anteriorly, these scales are very small, flake-like and they extend on the cleithral posterior process.

Posteriorly to the median fins, these scales are a much larger, irregular and scute-like shaped.

Between the nuchal horn and the dorsal fin, the dorsal ridge is composed by 7 to 9 scutes with a spiny

upper margin.

The ventral keel contains at least 4 scutes. The first three are located in the cloacal region. The posterior

one of these three has a spiny lower margin. A fourth spiny scute is associated with the ventral extremity

of the postcoelomic bone.

Joinvillichthys kriweti sp. nov.

um:lsid:zoobank.org:act:B8C4822F-44CA-4776-8455-452C5DBDCAlC

Figs 12-16

Diagnosis

Joinvillichthys with a body depth equal to 46.0 % of the standard length. A dorsal prominence present on

the frontal. Maxilla elongated. Farge parietal. Dermosupraoccipital sutured to the parietal and not to the

dermopterotic. Dermopterotic much longer than deep. Small dermosphenotic. Comma-shaped opercle.

Anterior ventral branch of the cleithrum lost. Broad and short pectoral spine articulated on the ventral

margin of the cleithral posterior process. Caudal fin double emarginated.

European Journal of Taxonomy 101: 1-27 (2014)

Etymology

The name of the new species is dedicated to Dr Jurgen Kriwet (Vienna) who has greatly improved our

knowledge of the pycnodontiform fishes.

Holotype

LEBANON: sample CLC S-137, a complete specimen from Haqel (Fig. 12), total length: 91 mm.

Paratype

LEBANON: sample AMNH 4517a (3698) and counterpart, an incomplete specimen from Hgula (Hay

1903: pi. 29, fig. 1); only the head and the beginning of the body are preserved, length: 63 mm.

Formation and locality

Marine Upper Cenomanian, Haqel and Hgula, Lebanon.

Morphometric data (Fig. 13)

The morphometric data are given in % of the holotype standard length (76 mm)

Length of the head (dermosupraoccipital included) . 54.2 %

Length of the cephalo-thorax (cleithrum included) . 75.3 %

Depth of the head (without the nuchal horn) . 43.8 %

Length of the nuchal horn . 27.7 %

Maximum depth of the body (just behind the nuchal horn) . 46.0 %

Fig. 12. Joinvillichthys kriweti gen. et sp. nov. Holotype, CLC S-137.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

Predorsal length. 16.6 %

Basal length of the dorsal fin . 9.8 %

Preanal length. 80.4 %

Basal length of the anal fin.6.4 %

Depth of the caudal peduncle.9.8 %

Osteology

1. The skull (Fig 14)

The general morphology of the skull is rather close to that of Joinvillichthys lindstroemi and the cranial

dermal bones also are ornamented with small tubercles. However, there are many small differences in

the head skeleton of the two fishes. Thus the description that follows will principally emphasize these

differences.

The skull is shorter and deeper than in Joinvillichthys lindstroemi. Its depth, from the upper margin of

the dermosupraoccipital to the lower margin of the cleithrum, is equal to 83 to 86 % of its length, from

the tip of the snout to the basis of the nuchal horn.

The rostrum lengthening is less pronounced. The prefrontal is broader and has a very sinuous suture with

the frontal. Its anterior tip also bears very small spines but is less outpacing of the lower jaw level. The

frontal is broader but does not outpace posteriorly the level of the orbit. The bone bears a small dorsal

prominence. The dermosupraoccipital is longer and is sutured to the parietal and the supratemporal but

not with the dermopterotic. The parietal is considerably larger. The dermopterotic is longer but much

thinner. The supratemporal is sutured to the parietal and reaches the dermopterotic at only one point. As

in Joinvillichthys lindstroemi , the long nuchal horn is supported only by the dermosupraoccipital.

Fig. 13. Joinvillichthys kriweti gen. et sp. nov. General reconstruction based on holotype, CLC S-137.

European Journal of Taxonomy 101: 1-27 (2014)

The orbitosphenoid and the pleurosphenoid are present in the orbit, just below the frontal, but the

basisphenoid is not visible.

The toothless premaxilla is longer and narrower. The toothless maxilla also is narrower and more

elongate. The lower jaw is composed with the same bones but is longer. The dentary bears two small

incisiform teeth and its ventral margin is denticulated. The articulation with the quadrate is located at the

level of the posterior border of the orbit.

A fragment of a large first infraorbital is preserved on the suture between the prefrontal and the premaxilla.

The sclerotic ring is visible in the orbit.

The hyomandibula and the preopercle are sutured together. The exposed part of the hyomandibula-

dermohyomandibula is larger than in Joinvillichthys lindstroemi but still much smaller than the

considerably enlarged preopercle. The opercle is broader and comma-shaped, with the sharp end dorsally

located. A part of the anterior ceratohyal and two small branchiostegal rays are visible behind the lower

jaw.

Fig. 14. Joinvillichthys kriweti gen. et sp. nov. Reconstruction of the skull and the pectoral girdle based

on holotype, CTC S-137.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

2. The girdles (Figs 14, 16)

The bones of the gigantic pectoral girdle have the same size and shape as in Joinvillichthys lindstroemi.

However, two important differences exist. The anterior ventral branch of the cleithrum is lost. No

postcleithrum is visible, but that is perhaps due to the fossilization. The pectoral spine is shorter, much

broader and is not articulated with the rear of the cleithrum but more anteriorly on its lower margin.

A small pelvic girdle is present. Indeed, a part of a vertically oriented pelvic bone is visible under a

broken part of the cleithrum.

3. The axial skeleton (Fig. 13)

The trunk is fusiform but proportionally deeper than in Joinvillichthys lindstroemi. The axial skeleton is

incomplete. Three vertebral segments are missing near the caudal peduncle. There are 16 neural spines

(the three missing ones included) before the epichordal series. Only 8 haemal spines are preserved. The

total number of haemal spines must be around 12 or 13. The neural and haemal spines are short but

broad. The neural and haemal arches surround almost completely the notochord. No ribs are visible. The

postcoelomic bone is well developed and backwardly oriented.

Fig. 15. Joinvillichthys kriweti gen. et sp. nov. Caudal skeleton of holotype, CLC S-137. The arrows

indicate the positions of the most external dorsal and ventral procurrent rays of the caudal fin.

European Journal of Taxonomy 101: 1-27 (2014)

4. The dorsal and anal fins (Fig. 13)

The short dorsal and anal fins are located at the mid-length of the body. There are 9 pterygiophores

and 9 rays in the dorsal fin. The anal fin contains 7 rays, but the number of anal pterygiophores is not

determinable. The first dorsal and anal ray is spiny. The other rays are segmented.

5. The caudal skeleton (Fig. 15)

The caudal skeleton of the holotype is partly preserved. There are 6 short and broad epichordals and 7

hypochordals. However, one or two anterior hypochordals are missing, so the complete series must be

composed of 8 or 9 elements. The fifth preserved hypochordal is strongly enlarged. No urodermal is

visible, but the region where theses bones are usually present is not preserved.

The caudal fin is double emarginated (Poyato-Ariza & Wenz 2002: fig. 36E) and contains 19 principal

segmented caudal rays, the 2 external being pointed and the 17 others branched. There are 6 ventral and

at least 4 dorsal procurrent rays.

2 mm a

Fig. 16. Joinvillichthys kriweti gen. et sp. nov. Holotype, CLC S-137. A. Dorsal ridge scutes. B. Anterior

flake-like body scales. C. Posterior scute-like body scale. D. Ventral keel scutes.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

6. Squamation (Fig. 16)

The squamation is the same as in Joinvillichthys lindstroemi. There are 7 spiny scutes in the dorsal

ridge and at least 3 spiny scutes in the ventral keel. The two posterior ventral scutes are associated with

the ventral margin of the postcoelomic bone. The body scales are slightly ornamented with tubercles.

Anteriorly, they are small and flake-like. Posteriorly, there are much larger, irregular and scute-like

shaped.

Pankowskichthys gen. nov.

um:lsid:zoobank.org:act:6C6C231C-40DC-4AD6-9250-2FD72E4EDClB

Type species: Pankowskichthys libanicus gen. et sp. nov. (by monotypy)

Diagnosis

As for the species (monospecific genus).

Etymology

The name of the new genus is dedicated to Mark Pankowski (Rockville, Maryland, U.S.A.), who

generously offered the holotype of Pankowskichthys libanicus gen. et sp. nov. to the Royal Belgian

Institute for Natural Sciences (IRSNB).

Pankowskichthys libanicus gen. et sp. nov.

um:lsid:zoobank.org:act:F3762EEA-A774-4377-9D4D-8E9DF3E28EE7

Figs 17-21

Diagnosis

Gladiopycnodontid with the elongate prefrontal forming a long, acuminate rostrum greatly outpacing

the lower jaw level. Fong frontal with a weakly developed dorsal prominence. Farge parietal. Small

dermopterotic. Orbitosphenoid present and separated from the mesethmoid. Premaxilla rather short,

toothless, with a hook-like anterior process, and sutured by its upper margin to the prefrontal. Small,

toothless triangular maxilla. Dentary curved and bearing 2 incisiform teeth. Hypertrophied trapezoid

preopercle covering the cheek. Exposed part of the hyomandibula-dermohyomandibula much smaller

than the preopercle. Extremely long nuchal horn, with a spiny posterior margin, and resting on both

the dermosupraoccipital and the parietal. Dermosphenotic narrow. Pectoral girdle closely associated

to the skull, forming a cephalo-thorax. Cleithrum hypertrophied, with a long, pointed, ventral branch.

Hypercleithrum hypertrophied. Small posttemporal. Very short and broad pectoral spine articulated on

the ventral margin of the cleithrum. Fong dorsal fin, beginning just behind the nuchal horn. Short anal

fin with 7 rays. A very broad hypural plate present in the hypochordal series. Body entirely covered by

large, scute-like scales. A short, ventral spine articulated on the postcoelomic bone.

Etymology

The species name refers to Febanon.

Holotype

FEBANON: sample IRSNB P 9278, a complete and well preserved specimen from Hgula (Fig. 17), total

length: 41 in in

Formation and locality

Marine Upper Cenomanian, Hgula, Febanon. The species has also been found at Haqel (Gayet et al.

2012: 87).

European Journal of Taxonomy 101: 1-27 (2014)

Fig. 17. Pankowskichthys libanicus gen. et sp. nov. Holotype, IRSNB P 9278.

Fig. 18. Pankowskichthys libanicus gen. et sp. nov. General reconstruction based on holotype, IRSNB

P 9278.

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

Morphometric data (Fig. 18)

The morphometric data are given in % of the standard length of the holotype (36 mm).

Length of the head (dermosupraoccipital and parietal included) . 73.1 %

Length of the cephalo-thorax (cleithrum included) . 72.5 %

Depth of the head (without the nuchal horn). 42.4 %

Length of the nuchal horn. 83.5 %

Maximum depth of the body (just behind the nuchal horn).35.4 %

Predorsal length. 82.6 %

Basal length of the dorsal fin. 20.3 %

Preanal length. 75.9 %

Basal length of the anal fin.8.9 %

Depth of the caudal peduncle.8.5 %

Osteology

1. The skull (Fig 19)

The head and the pectoral girdle are closely associated, forming a cephalo-thorax that is gigantic when

compared to the body size. This character and the feeble ossification of the axial skeleton probably

indicate that the concerned sample is a juvenile fish. The dermal bones are ornamented with tubercles,

alveoli and thin ridges.

Fig. 19. Pankowskichthys libanicus gen. et sp. nov. Reconstruction of the skull and the pectoral girdle

based on holotype, IRSNB P 9278.

European Journal of Taxonomy 101: 1-27 (2014)

The rostrum is longer than in Joinvillichthys gen. nov. It is formed by the acuminate anterior extremity

of the prefrontal. This rostrum greatly outpaces the lower jaw level. The preorbital length, rostrum

included, represents 54.5 % of the total length of the head. The prefrontal is long, anteriorly pointed and

rather broad posteriorly, hiding completely the vomer and a great part of the mesethmoid.

The frontal is narrow, rather long and exhibits a weakly developed median protuberance located

just above the orbit. The elongate posterior part of the frontal extends far behind the orbit, under

the dermosupraoccipital ventral margin, and reaches the parietal. The dermopterotic is small but the

dermosupraoccipital and the parietal are large bones that protrude posteriorly. The small autosphenotic

is located in front of the dermopterotic and just below the posterior extremity of the frontal. An

extremely long nuchal horn with a spiny posterior margin is articulated on both the parietal and the

dermosupraoccipital.

The parasphenoid is very long, straight and toothless. Its posterior extremity greatly outpaces the rear

of the skull. The orbitosphenoid is separated from the mesethmoid. No other endocranial bone of the

braincase and no bone of the palato-quadrate arch are visible.

The premaxilla and the maxilla are toothless. There is no supramaxilla. The premaxilla is long but

much shorter than the prefrontal, to which it is sutured by its upper margin. This premaxilla exhibits

a small, anterior hook-shaped process, a broad anterior part and a much narrower posterior region.

The premaxilla does not contribute to the rostrum. The small, triangular maxilla is located below the

posterior part of the premaxilla. The dentary, reduced to its ventral branch, is well developed. It bears

two very small, incisiform teeth. Its posterior part forms a right angle with its anterior extremity. Its

lower margin is spiny. The angular is a rather large bone. Only a very small part of the prearticular is

visible. The articular is not preserved.

The orbit is wide. The long and very thin dermosphenotic is placed against the autosphenotic and the

dermopterotic. No other bone of the orbital series is present.

HYCO

Fig. 20. Pankowskichthys libanicus gen. et sp. nov. Caudal skeleton of holotype, IRSNB P 9278.

TAVERNE L. & CAPASSO F., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

The hyomandibula and the preopercle are sutured together. The exposed part of the hyomandibula-

dermohyomandibula is deeper than broad but much smaller than the hypertrophied, trapezoid-shaped

preopercle that covers the cheek. The opercle is a long, rod-like bone pressed between the preopercle

and the pectoral girdle.

2. The girdles (Figs 18-19)

The hypertrophied pectoral girdle is pressed against the skull. The bones are ornamented with tubercles,

alveoli and thin ridges. The gigantic cleithrum possesses a long, narrow and pointed ventral branch, with

a serrated lower margin, and a very broad posterior process. The hypercleithrum (= supracleithrum) is

a large bone, broader ventrally than dorsally. A small, triangular posttemporal, with an acuminate upper

extremity, is located near the parietal. A small postcleithrum is visible behind the hypercleithrum. The

pectoral fin is replaced by a short but very broad spine that is articulated and partly fused to the ventral

margin of the wide cleithral posterior process.

A fragment of a pelvic ray is visible a little before the postcoelomic bone. The pelvic bones are not

preserved.

3. The axial skeleton (Fig. 18)

The axial skeleton is poorly ossified. Only the first thirteen vertebral segments are partly preserved. The

complete vertebral axis probably contained a little less than twenty segments. Well developed neural

arches are present but the haemal arches are not visible. Thus, the notochord is not surrounded by the

arches. The first neural and haemal spines are long and the last ones very short. Fragments of ribs are

visible. The postcoelomic bone is broad but rather short.

4. The dorsal and anal fins (Fig. 18)

The dorsal fin has a rather long basis and begins just behind the nuchal horn. The number of rays and

pterygiophores is unknown. The anal fin has a shorter basis and contains 7 rays. Traces of at least 5 anal

pterygiophores are visible.

5. The caudal skeleton (Fig. 20)

The caudal skeleton is incompletely preserved. Only the hypochordal series is present with 5 elements.

The fourth hypochordal is greatly enlarged, forming a very broad hypural plate. This plate probably

results from the fusion of several hypochordals. The epichordals and the urodermals are not known.

Only a part of the caudal fin is present.

0.5 mm

Fig. 21. Pankowskichthys libanicus gen. et sp. nov. A scute-like body scale of holotype, IRSNB P 9278.

European Journal of Taxonomy 101: 1-27 (2014)

6. Squamation (Fig. 21)

The squamation is badly preserved. However, some fragments and prints of scales are visible on the

entire body. These scales are large, scute-like, irregularly shaped and ornamented with tubercles and

alveoli.

A small spine is articulated on the ventral extremity of the postcoelomic bone.

Discussion

1. Joinvillichthys gen. nov. and Pankowskichthys gen. nov. within Neopterygii

Joinvillichthys gen. nov. and Pankowskichthys gen. nov. share a few very peculiar characters: (1) there

is a dermosupraoccipital, (2) the maxilla is plate-like and toothless, (3) the supramaxillae are lost, (4)

the mesethmoid is very long and includes the lateral ethmoids, (5) a dermohyomandibula is fused with

the hyomandibula, (6) the hyomandibula is sutured to the preopercle, (7) the preopercle is greatly

hypertrophied, trapezoid in shape and it covers a great part of the cheek, (8) the opercle is severely

reduced, (9) the subopercle and interopercle are lost, (10) the parasphenoid is elongate and toothless,

(11) the pelvic girdle is reduced, (12) a postcoelomic bone is present and (13) the neural and haemal

arches do not fuse together. Within Neopterygii, the association of these thirteen characters states is

found only in the Pycnodontomorpha, a superorder recently erected by Nursall (2010) for the former

Pycnodontiformes and to which the two new genera can be allocated.

Nursall (2010) divided the Pycnodontomorpha into two orders, the Gyrodontiformes for the families

Mesturidae and Gyrodontidae and the Pycnodontiformes as a new usage for the former Pycnodontoidei.

Species belonging to the Gyrodontiformes have two dermosupraoccipitals, a dentary with dorsal and

ventral branches, styliform teeth on the premaxilla and the dentary, small bony tesserae covering the

snout, the cheek and the gular region, while they also lack tubular infraorbitals. Species belonging to

the Pycnodontiformes possess only one dermosupraoccipital, a dentary reduced to its ventral branch,

incisiform teeth on the premaxilla and the dentary and true infraorbitals even when a few bony tesserae

are preserved on the cheek or in the gular region. Thus, the osteological characters of Joinvillichthys

gen. nov. and Pankowskichthys gen. nov. refer these two fish genera to the Pycnodontiformes and not to

the Gyrodontiformes.

The two new Lebanese genera present three remarkable apomorphies. The prefrontal and the premaxilla

combined or the prefrontal alone form a long rostrum that extends more or less beyond the lower jaw

level. The upper margin of the toothless premaxilla is sutured to the lower margin of the elongate and

broadened prefrontal. The pectoral fin is lost and replaced by a strong pectoral spine articulated on the

cleithrum. These three characters only exist in the family Gladiopycnodontidae (Taverne & Capasso

2013) and not in any other pycnodontiform lineage.

2. Joinvillichthys gen. nov. and Coccodus

As was stated above, Joinvillichthys lindstroemi was originally ranged by Davis (1890) in Coccodus ,

a Late Cretaceous specialized pycnodontiform genus from Lebanon. This generic attribution was

explicitely contested by Poyato-Ariza & Wenz (2002: 145). A short comparison between the two

concerned genera is thus useful. The 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 (Taverne & Capasso 2014).

Coccodus exhibits a normal pycnodontiform snout, with a long, toothed premaxilla overlying a

narrow prefrontal. There is no rostrum, the upper and lower jaws being located at the same level. The

dermosupraoccipital bears a large horn and develops a short posterior process, but there is no nuchal

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

horn. The teeth on the vomer and the prearticular are typically pycnodontiform, i. e. , large, molariform

and ranged in regular rows. The preopercle is not very wide and does not cover completely the ventral

branch of the hyomandibula. There is a normal pectoral fin with soft rays. Such a skeleton greatly differs

from that of Joinvillichthys lindstroemi.

These important cranial and pectoral differences clearly show that Joinvillichthys lindstroemi cannot be

reported to Coccodus.

3. The validity of Joinvillichthys kriweti gen. et sp. nov.

Joinvillichthys kriweti gen. et sp. nov. is similar to J. lindstroemi. However, the two species differ in

many morphological and osteological details. Joinvillichthys kriweti gen. et sp. nov. has a deeper head

and a deeper body. Many bones have different shapes and sizes in the two species. This is the case, for

example, for the frontal, the parietal, the dermosupraoccipital, the dermopterotic, the dermosphenotic,

the premaxilla, the maxilla, the preopercle, the opercle, the exposed part of the hyomandibula-

dermohyomandibula, the cleithrum and the pectoral spine. Joinvillichthys kriweti gen. et sp. nov. also

exhibits a double emarginated caudal fin, whereas J. lindstroemi has a caudal fin with a convex posterior

margin. These differences amply justify the erection of the new species, Joinvillichthys kriweti gen. et

sp. nov.

4. The validity of Pankowskichthys gen. nov.

Pankowskichthys gen. nov. and Joinvillichthys gen. nov. possess the same basal anatomy but they greatly

differ in the details of their skeleton. Almost all the bones of the cranium, the jaws and the pectoral girdle

of the two fishes exhibit different shapes and sizes.

Some differences between the two genera could be owing to allometry and ontogenetic growth, for

instance, the length of the nuchal horn, the respective proportions of the cephalo-thorax and body, and

the development of the neural and haemal arches.

Other osteological differences, however, are not allometric:

(1) The frontal is sutured with the dermopterotic in Pankowskichthys gen. nov. That is not the case in

Joinvillichthys gen. nov.

(2) The premaxilla of Pankowskichthys gen. nov. bears a well developed, anterior hook-like process.

Such a process is absent on the premaxilla of Joinvillichthys gen. nov..

(3) Joinvillichthys gen. nov. has a short dorsal fin located in the middle of the back, far from the nuchal

horn. Pankowskichthys gen. nov. has a longer dorsal fin reaching the nuchal horn.

(4) A series of dorsal ridge scutes is present in Joinvillichthys gen. nov., but these are absent in

Pankowskichthys gen. nov.

(5) A greatly hypertrophied hypural plate is present in the caudal skeleton of Pankowskichthys gen.

nov. In Joinvillichthys gen. nov. and in the other gladiopycnodontid genera, some hypochordals are

broadened but never hypertrophied.

(6) The body is entirely covered by scute-like scales in Pankowskichthys gen. nov. In Joinvillichthys

gen. nov., the greatest part of the body is covered by flake-like scales and the scute-like scales are

only present in the tail region.

Pankowskichthys gen. nov. also shares with the more specialized gladiopycnodontid genera a series of

apomorphies not present in Joinvillichthys gen. nov.:

(1) An anal spine is associated with the postcoelomic bone. This spine is short in Pankowskichthys

gen. nov. and in one still undescribed genus of the family (pers. obs.). This spine is very large

in Rostropycnodus and in another undescribed genus (pers. obs.), and becomes really gigantic

in Gladiopycnodus (Taverne & Capasso 2013: figs 2, 18, 20). In Joinvillichthys gen. nov., as in

Monocerichthys, the most primitive genus of the family, only postcloacal scales are present at the

European Journal of Taxonomy 101: 1-27 (2014)

level of the postcoelomic bone, as normally in pycnodontid fishes (Taverne & Capasso 2013: figs 8,

12 ).

(2) The pectoral spine is shortened but considerably broadened and more or less fused with the

cleithrum in Pankowskichthys gen. nov. and in a few other specialized genera (Taverne & Capasso

2013: fig. 19). In Joinvillichthys gen. nov., the pectoral spine is longer, thinner and simply fixed to

the cleithrum.

(3) The rostrum is elongated and becomes acuminate in Pankowskichthys gen. nov. and the more evolved

genera of the family, the lower jaw remaining rather short. This elongation reaches its maximum

in Gladiopycnodus and Rostropycnodus (Taverne & Capasso 2013: figs 3, 19). In Joinvillichthys

gen. nov., as in Monocerichthys , the rostrum outpaces the lower jaw level only slightly (Taverne &

Capasso 2013: fig. 9).

(4) The nuchal horn is fixed to the dermosupraoccipital and the parietal in Pankowskichthys gen. nov.

and in the more specialized genera having retained a nuchal horn (pers. obs.). In Joinvillichthys gen.

nov., as in Monocerichthys , the parietal is not involved in the cranial articulation of the nuchal horn

(Taverne & Capasso 2013: fig. 9).

Such a number of differences, and especially the apomorphies shared with the more evolved genera

of the family, makes it unreasonable to consider Pankowskichthys gen. nov. as a juvenile specimen of

Joinvillichthys gen. nov. and justifies the creation of a unique new genus for the former fish.

5. Joinvillichthys gen. nov. and Pankowskichthys gen. nov. within Gladiopycnodontidae

Joinvillichthys gen. nov. and Pankowskichthys gen. nov. are rather primitive gladiopycnodontid fishes.

Indeed, they preserved the nuchal horn that was lost in the more evolved members of the family (Taverne

& Capasso 2013: figs 3, 19) and they had not acquired the frontal horn and the strong anal spine present

in some advanced genera (Taverne & Capasso 2013: figs 2, 4, 18-20).

The skull and pectoral girdle of Joinvillichthys gen. nov. are not very different from those of

Monocerichthys. The rostrum is a little more elongate and more acuminate in the new genus and its

nuchal horn is a little longer and rests only on the dermosupraoccipital, with no implication of the

supratemporal or posttemporal. Joinvillichthys gen. nov. also possesses dorsal ridge and ventral keel

scutes, in opposition to Monocerichthys that lacks these structures (Taverne & Capasso 2013: fig. 8).

The presence of scute-like scales in the caudal region of Joinvillichthys gen. nov. is another apomorphic

character absent in Monocerichthys , in which only small, flake-like scales cover the entire body (Taverne

& Capasso 2013: fig. 14).

Pankowskichthys gen. nov. is more apomorphic than Monocerichthys and Joinvillichthys gen. nov., as

seen in section 4 of the present discussion.

The phylogeny within the Gladiopycnodontidae will be studied in a forthcoming paper, which includes

the description of four other genera of this family (Taverne & Capasso, in prep.).

Acknowledgments

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

delTAbruzzo (Chieti, Italy) for allowing us to study the fossil fishes of Tuigi Capasso’s collection.

We are grateful to M. Adriano Vandersypen and M. Wilfried Miseur of the Royal Belgian Institute of

Natural Sciences (Brussels, Belgium) and to M. Tuciano Tullo from the University of Chieti (Chieti,

Italy) for their technical help. We also thank Dr Thomas Mors, senior curator of Palaeontology at the

Swedish Museum of Natural History (Stockholm, Sweden), Dr John Maisey from the Department of

Palaeontology of the American Museum of Natural History (New York, USA) and Dr Etienne Steurbaut,

head of the former Department of Palaeontology at the Royal Belgian Institute of Natural Sciences

TAVERNE L. & CAPASSO L., The “ Coccodus ” lindstroemi species complex (Pycnodontiformes)

(Brussels, Belgium), for allowing access to their collections and for providing photos of some samples.

We are also indebted to the two anonymous reviewers who have read and commented on our text.

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genres. Palaeontos 25: 3—43.

Manuscript received: 26 February 2014

Manuscript accepted: 8 September 2014

Published on: 5 November 2014

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 consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, De nm ark