A new genus and species of pycnodontid fish Flagellipinna rhomboides, gen. et sp. nov. (Neopterygii, Pycnodontiformes), from the Upper Cretaceous (Cenomanian) of Lebanon, with notes on juvenile form and ecology

Figures & data

FIGURE 1. Specimens of Flagellipinna rhomboides, gen. et sp. nov. A, MNHN.F.HAK2003, holotype. B, MNHN.F.HAK2001, paratype, possible late juvenile/subadult stage. C, MNHN.F.HAK1972a, paratype, part. D, MNHN.F.HAK1972b, paratype, counterpart. Scale bars equal 1 cm.

FIGURE 2. Flagellipinna rhomboides, gen. et sp. nov., reconstruction of the skeleton based on MNHN.F.HAK1972a, MNHN.F.HAK1972b, MNHN.F.HAK2001, and MNHN.F.HAK2003. Anatomical characters based on the holotype and the paratypes; skull roof restored from the holotype, whereas the lower jaw is based on both the holotype and MNHN.F.HAK2001; pectoral girdle is intermediate between the conditions seen in the holotype and MNHN.F.HAK2001; dorsal ridge scale series based mainly on MNHN.F.HAK1972b; caudal endoskeleton based on a combination of MNHN.F.HAK1972b and specimen figured on page 93 of Gayet et al. (2012); arcocenta based on paratype MNHN.F.HAK1972b. Reconstruction of paired fins and the caudal fin is hypothetical because they are poorly preserved in all specimens. Caudal endoskeleton that is not preserved in any specimen is represented by hypothethical bones shown here in gray. Scale bar equals 1 cm.

FIGURE 3. Flagellipinna rhomboides, gen. et sp. nov., MNHN.F.HAK2003, holotype, skull and pectoral girdle. A, photograph of the head and anterior region of the body. Photo by J.J.C. B, camera lucida drawing of the skull bones seen in A. Abbreviations: 1drs, first dorsal ridge scale; ang, angular; art, articular; br, branchiostegal rays; cha, anterior ceratohyal; chp, posterior ceratohyal; cl, cleithrum; den, dentalosplenial; dhyo, dermohyomandibular; dps, dermopterosphenotic; dspo, dermosupraoccipital; ect, ectopterygoid; ent, entopterygoid; hh, hypohyal; ih, interhyal; mes, mesethmoid; met, metapterygoid; mx, maxilla; op, operculum; par, parietal; pmx, premaxilla; pop, preoperculum; pp, postparietal; ppr, postparietal process; prart, prearticular; prep, preparietal; psph, parasphenoid; q, quadrate; scl, sclerotic ring; vom, vomer. Scale bar equals 1 cm.

FIGURE 4. Flagellipinna rhomboides, gen. et sp. nov., MNHN.F.HAK2001, skull and pectoral girdle of juvenile/subadult specimen. Dotted lines indicate where sutures between bones could be visible but are difficult to confirm fully in the specimen. Ossification is poor in the skull roof, so arrows are used frequently to show what parts of fragmented material could belong to certain bones. Asterisk used to show displaced vomerine tooth overlapping ventral margin of parasphenoid. A, photograph of the head and anterior region of the body. Photo by J.J.C. B, camera lucida drawing of the skull bones seen in A. Abbreviations: 1drs, first dorsal ridge scale; ang, angular; art, articular; br, branchiostegal rays; cha, anterior ceratohyal; chp, posterior ceratohyal; cl, cleithrum; cpr, coronoid process; den, dentalosplenial; dhyo, dermohyomandibular; dps, dermopterosphenotic; dspo, dermosupraoccipital; ect, ectopterygoid; ent, entopterygoid; ih, interhyal; mes, mesethmoid; met, metapterygoid; mx, maxilla; op, operculum; par, parietal; pmx, premaxilla; pop, preoperculum; pp, postparietal; prep, preparietal; psph, parasphenoid; q, quadrate; scl, sclerotic ring. Scale bar equals 1 cm.

FIGURE 5. Flagellipinna rhomboides, gen. et sp. nov., MNHN.F.HAK1972b, paratype, infraorbital series. Scale bar equals 1 cm.

FIGURE 6. Flagellipinna rhomboides, gen. et sp. nov., MNHN.F.HAK2003, holotype, remains of pelvic girdle. Scale bar equals 1 cm.

FIGURE 7. Flagellipinna rhomboides, gen. et sp. nov., MNHN.F.HAK1972b, paratype, caudal endoskeleton. Abbreviations: hypo 1–5, hypochordals 1–5; phyp, parhypural. Scale bar equals 1 cm.

FIGURE 8. Flagellipinna rhomboides, gen. et sp. nov. Reconstruction of the ventral ridge scales and cloaca based on the holotype MNHN.F.HAK2003. Abbreviations: cs, cloacal scales; pcb, post coelomic bone; pvrs, postcloacal ventral ridge scale; vrs, ventral ridge scales. Scale bar equals 1 cm.

FIGURE 9. Flagellipinna rhomboides, gen. et sp. nov., MNHN.F.HAK1972b, paratype, dorsal ridge scales. Scale bar equals 1 cm.

TABLE 1. Relevant measurements (in cm), proportions, and meristic characters of the articulated specimens of the type series of Flagellipinna rhomboides, gen. et sp. nov.

Dimension	Holotype	MNHN.F.HAK2001	MNHN.F.HAK1972
SL	8.23	7.04	9.72
PDOR	6.28	5.33	3.47
%SL	76.30	75.70	35.70
PANA	4.31	4.60	—
%SL	52.40	65.30	—
PPEC	2.92	2.87	—
%SL	35.50	40.80	—
HD	4.09	5.17	—
%SL	49.70	73.40	—
MBD	8.02	10.07	—
%SL	97.40	143.00	—
%HD/MBD	51.00	51.30	—
HL	4	3.54	—
%SL	48.60	50.30	—
DFH	2.05	1.12	4.39
%SL	24.90	15.90	45.20
%HD/HL	103.30	146.00	—
DP	(36)	38	46
AP	(31)	(38)	46
VE	(22)	27	(24)
AE	7	5	—
AS	6	4	—
DRS	10	(17)	(24)
VRS	12	14	—
PCL	1	2	—
ACS	1	1	—
PCS	1	—	—
°APROF	47.60	65.79	58.51

Parentheses indicate that the corresponding measure is an estimate or that the figure is based on a poorly preserved structure. All SLs are estimated measurements due to incomplete preservation in all specimens. Abbreviations: °APROF, slope of anterior profile; %HD/HL, percentage of ratio between head depth and head length; %HD/MBD, percentage of ratio between head depth and maximum body length; %SL, percentage of ratio between standard length and above measurement; ACS, number of anterior modified cloacal scales; AE, number of abdominal vertebrae (epaxial elements); AP, number of anal pterygiophores; AS, number of anterior autogenous neural spines; DFH, dorsal fin height; DP, number of dorsal pterygiophores; DRS, number of dorsal ridge scales; HD, head depth (from highest point of dermosupraoccipital to lowest point of cleithrum); HL, head length (from anterior border of premaxillary to posterior border of cleithrum); MBD, maximum body depth; PANA, preanal distance; PCL, number of postcloacal ventral ridge scales; PCS, number of posterior modified cloacal scales; PDOR, predorsal distance; PPEC, prepectoral distance; SL, standard length; VE, number of vertebrae (epaxial elements excluding those supporting precurrent and principal caudal fin rays); VRS, total number of ventral ridge scales.

FIGURE 10. Pycnodont taxa that share prognathism with Flagellipinna rhomboides, gen. et sp. nov. A, Iemanja palma, NSM PV-20384. B, Anomoeodus nursalli, IPFUB-Uña Pyc 1. Scale bars equal 10 cm (A) and 1 cm (B).

APPENDIX 1. List of all complete pycnodont taxa found in the Lebanese Cenomanian deposits.

Taxon	Family	Location
Acrorhinichthys poyatoi Taverne and Capasso, 2015b	Pycnodontidae	Ein Nammoura, Haqel
Akromystax tilmachiton Poyato-Ariza and Wenz, 2005	Pycnodontidae	Haqel
Coccodus armatus Pictet, 1850	Coccodontidae	Haqel
Coccodus insignis Hay, 1903	Coccodontidae	Hjoûla, Haqel
Corusichthys megacephalus Taverne and Capasso, 2014b	Coccodontidae	Haqel
Ducrotayichthys cornutus Taverne and Capasso, 2015a	Gladiopycnodontidae	Haqel
Gebrayelichthys uyenoi Nursall and Capasso, 2004	Gebrayelichthyidae	Hjoûla, Haqel
Gebrayelichthys vexillarius Nursall and Capasso, 2004	Gebrayelichthyidae	Haqel
Gladiopycnodus byrnei Marramà et al., 2016	Gladiopycnodontidae	Hjoûla
Gladiopycnodus karami Taverne and Capasso, 2013a	Gladiopycnodontidae	Haqel
Haqelpycnodus picteti Taverne and Capasso, 2018a	Pycnodontidae	Haqel
Hayolperichthys pectospinus Taverne and Capasso, 2015a	Gladiopycnodontidae	Haqel
Hensodon spinosus Hennig, 1907	Coccodontidae	Haqel
Ichthyoceros spinosus Gayet, 1984	Gladiopycnodontidae	Haqel
Joinvillichthys kriweti Taverne and Capasso, 2014a	Gladiopycnodontidae	Hjoûla, Haqel
Joinvillichthys lindstroemi Davis, 1890	Gladiopycnodontidae	Hjoûla, Haqel
Libanopycnodus wenzi Taverne and Capasso, 2018b	Pycnodontidae	Ein Nammoura
Maraldichthys verticalis Taverne and Capasso, 2014c	Gebrayelichthyidae	Haqel
Monocerichthys scheuchzeri Taverne and Capasso, 2013a	Gladiopycnodontidae	Ein Nammoura, Hjoûla
‘Nursalia’ goedeli Heckel, 1854	Pycnodontidae	Haqel
Nursallia tethysensis Capasso et al., 2009	Pycnodontidae	Ein Nammoura, Hjoûla
Palaeobalistum sp.	Pycnodontidae	Hjoûla, Haqel
Pankowskichthys libanicus Taverne and Capasso, 2014a	Gladiopycnodontidae	Ein Nammoura, Haqel
Paracoccodus woodwardi Taverne and Capasso, 2014b	Coccodontidae	Haqel
Proscinetes, sp. indet. Forey et al., 2003	Pycnodontidae	Ein Nammoura
Rhinopycnodus gabriellae Taverne and Capasso, 2013b	Pycnodontidae	Haqel
Rostropycnodus gayeti Taverne and Capasso, 2013a	Gladiopycnodontidae	Haqel
Sigmapycnodus giganteus Taverne and Capasso, 2018b	Pycnodontidae	Haqel
Stenoprotome hamata Hay, 1903	Gladiopycnodontidae	Haqel
Trewavasia carinata Davis, 1887	Coccodontidae	Haqel
Tricerichthys wenzi Taverne and Capasso, 2015a	Gladiopycnodontidae	Haqel

Gayet, M., P. Abi Saad, and O. Gaudant. 2012. Les fossiles du Liban: Memoire du Temps. Éditions Desiris, Meolan-Revel, France, 184 pp.

 Google Scholar

Taverne, L., and L. Capasso. 2015b. Osteology and relationships of Acrorhinichthys poyatoi gen. et sp. nov. (Pycnodontiformes) from the marine Late Cretaceous of Lebanon. European Journal of Taxonomy 116:1–30.

 Web of Science ®Google Scholar

Poyato-Ariza, F. J., and S. Wenz. 2005. Akromystax tilmachiton gen. et sp. nov., a new pycnodontid fish from the Lebanese Late Cretaceous of Haqel and En Nammoura. Journal of Vertebrate Paleontology 25:27–45. doi: 10.1671/0272-4634(2005)025[0027:ATGESN]2.0.CO;2

 Web of Science ®Google Scholar

Pictet, F. J. 1850. Description de quelques poissons fossiles du Mont Liban. Imprimerie De Jules-Guillaume Fick, Geneva, 59 pp.

 Google Scholar

Hay, O. P. 1903. On a collection of Upper Cretaceous fishes from Mount Lebanon, Syria, with descriptions of four new genera and nineteen new species. Bulletin of the American Museum of Natural History 19:395–452.

 Google Scholar

Taverne, L., and L. Capasso. 2014b. Osteologie et phylogenie des Coccodontidae, une famille remarquable de poissons Pycnodontiformes du Cretace superieur marin du Liban, avec la description de deux nouveaux genres. Palaeontos 25:3–43.

 Google Scholar

Taverne, L., and L. Capasso. 2015a. New data on the osteology and phylogeny of Gladiopycnodontidae (Pycnodontiformes), a tropical fossil fish family from the marine Upper Cretaceous of Lebanon, with the description of four genera. Geo-Eco-Trop 39:217–246.

 Google Scholar

Nursall, J. R., and L. Capasso. 2004. Gebrayelichthys (novum), an extraordinary genus of neopterygian fishes from the Cenomanian of Lebanon; pp 317–340 in G. Arratia and A. Tintori (eds.), Mesozoic Fishes 3: Systematics, Paleoenvironments and Biodiversity. Dr. F. Pfeil, Munich, Germany.

 Google Scholar

Marramà, G., B. Villier, F. M. Dalla Vecchia, and G. Carnevale. 2016. A new species of Gladiopycnodus (Coccodontoidea, Pycnodontomorpha) from the Cretaceous of Lebanon provides new insights about the morphological diversification of pycnodont fishes through time. Cretaceous Research 61:34–43. doi: 10.1016/j.cretres.2015.12.022

 Web of Science ®Google Scholar

Taverne, L., and L. Capasso. 2013a. 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.

 Web of Science ®Google Scholar

Taverne, L., and L. Capasso. 2018a. Osteology and phylogenetic relationships of Haqelpycnodus picteti gen. and sp. nov., a new pycnodont fish genus (Pycnodontidae) from the marine Late Cretaceous tropical sea of Lebanon. Geo-Eco-Trop 42:117–132.

 Google Scholar

Hennig, E. 1907. Ueber einige Pyknodonten vom Libanon. Centralblatt für Mineralogie, Geologie und Palâontologie 1907:360–371.

 Google Scholar

Gayet, M. 1984. Ichthyoceros spinosus nov. gen., nov. sp., du Cénomanien inférieur de Hakel (Liban) et ses affinités avec le genre Trewavasia (Pisces, Pycnodontiformes, Coccodontidae). Bulletin du Muséum national d’Histoire naturelle 3:287–307.

 Google Scholar

Taverne, L., and L. Capasso. 2014a. 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.

 Web of Science ®Google Scholar

Davis, J. W. 1890. On a new species of Coccodus (C. lindstroemi, Davis). The Quarterly Journal of the Geological Society of London 46:565–568. doi: 10.1144/GSL.JGS.1890.046.01-04.36

 Google Scholar

Taverne, L., and L. Capasso. 2018b. Osteology and relationships of Libanopycnodus wenzi gen. et sp. nov. and Sigmapycnodus giganteus gen. et sp. nov. (Pycnodontiformes) from the Late Cretaceous of Lebanon. European Journal of Taxonomy 420:1–29.

 Web of Science ®Google Scholar

Taverne, L., and L. Capasso. 2014c. Osteologie et relations phylogenetiques des Gebrayelichthyidae (Halecostomi, Pycnodontomorpha), une extraordinaire famille de poissons du Cretacee superieur marin du Liban, avec la description d’unnouveau genre. Palaeontos 25:44–68.

 Google Scholar

Heckel, J. 1854. Über den Bau und die Eintheilung der Pycnodonten, nebst kurzer Beschreibung einiger neuen Arten derselben. Sitzungsberichte der kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse 12:433–464.

 Google Scholar

Capasso, L. L., P. Abi Saad, and L. Taverne. 2009. Nursallia tethysensis sp. nov., a new pycnodont fish (Neopterygii: †Halecostomi) from the Cenomanian of Lebanon. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 79:117–136.

 Google Scholar

Forey, P. L., L. Yi, C. Patterson, and C. E. Davies. 2003. Fossil fishes from the Cenomanian (Upper Cetaceous) of Namoura, Lebanon. Journal of Systematic Palaeontology 1:227–330. doi: 10.1017/S147720190300107X

 Web of Science ®Google Scholar

Taverne, L., and L. Capasso. 2013b. Osteology and relationships of Rhinopycnodus gabriellae gen. et sp. nov. (Pycnodontiformes) from the marine Late Cretaceous of Lebanon. European Journal of Taxonomy 67:1–14.

 Web of Science ®Google Scholar

Davis, J. W. 1887. The fossil fishes of the Chalk of Mount Lebanon in Syria. Scientific Transactions of the Royal Dublin Society 3:457–636.

 Google Scholar