Vol. 3 No. 2 page 33-37 DOI: 10.14456/randk.2017.22 RESEARCH & KNOWLEDGE Research Article Revision of the batomorphs from the of Lebanon, and their impact on our understanding of the early step of the evolution of the clade

Georges Kachacha1, *, Gilles Cuny2, Dany Azar3 and Pierre Abi Saad4

1Department of EvoGenomics, Natural History Museum of Denmark, Copenhagen, Denmark 2UMR CNRS 5276 ENS LGLTPE, Université Claude Bernard Lyon 1, Université de Lyon, Villeurbanne Cedex, France 3 Natural Sciences Department, Faculty of Sciences 2, Lebanese University, Fanar, Lebanon. 4 Memory of Time, Byblos, Lebanon

(Received 20 May 2017; accepted 22 June 2017)

Abstract - Lebanon is endowed with its outstanding preservation lagerstätten of fossil fish from the Upper Cretaceous. The batomorphs are represented by 16 of belonging to 9 genera and 4 families however, their phylogeny remains poorly understood. Also, their diversity is possibly underestimated, compared to the great diversification event observed in the Upper Cretaceous, by lumping the majority of Lebanese rhinobatids in the genus Rhinobatos, whereas their relationships with the modern species of this genus are unclear. We discuss herein the attribution of three lebanese -like “Rhinobatids” species included into two new genera to Rhynchobatidae. These species together with Rhynchobatus possess a combination of primitive characters such as propterygium failing to reach the level of the nasal capsules and pectoral fin posterior corner not reaching the level of the pelvic fin. Three additional characters are supporting this clade in the present analysis together with two plesiomorphies. The remaining present a closer relationship with Rajidae than to Pristidae and so the order recently erected by Naylor et al., (2012) is not recovered in our analysis.

Keywords: Batomorpha, Upper Cretaceous, Rhinobatidae, cladistics

1. Introduction and several Rhinobatoids) in the The Superorder Batomorpha (skates, stingrays and their phylogenetic tree of batomorphs and within Rajiformes is allies) presents the largest taxonomic diversity among far from clear. extant (≈630 of ≈1221 species) (Compagno, In the Upper Cretaceous of Lebanon, the batomorphs 2005; Fowler, 2005; Naylor et al., 2012). It appeared are represented by 16 species of Rajiformes belonging to during the (Underwood, 2006) and is divided into 9 genera and 4 families. The Rajiformes phylogeny remains three orders: Rajiformes, Myliobatiformes and Torpedini- poorly understood with the “Rhinobatidae” forming a formes (Cappetta, 2012). This cartilaginous fish lineage is polyphyletic group (Brito and Dutheil, 2004; Brito et al, recognized as a monophyletic group sister to all living 2013; Claeson et al. 2013) and the position of the Sclero- (Selachimorpha), based on both molecular and rhynchoidei fluctuating due to the lack of available morphological data (Douady et al., 2003; Maisey et al., well-preserved specimens. Similarly, the phylogenetic 2004; Winchell et al., 2004; Aschliman et al., 2012a; affinity of the Lebanese fossil Rajidae and Cylobatidae is Naylor et al, 2012; Carvalho, 1996). Nonetheless, the not yet solved. The diversity of the Lebanese Rajiformes interrelationships within batomorphs remain controversial is possibly underestimated, compared to the great diversi- amongst morphologists and molecular biologists, with the fication event observed in the Upper Cretaceous (Guinot most contentious issues concerning the most basal group and Cavin, 2015), by lumping the majority of Lebanese position. The phylogenetic relationships within batomorphs rhinobatids in the genus Rhinobatos, whereas their relation- are yet even more complicated when fossils are considered ships with the modern species of this genus are unclear because their fossil record is mostly restricted to teeth, (Cappetta, 2012; Claeson et al, 2013; Brito et al., 2013). dermal denticles and occasionally, fin spines and calcified The outstanding preservation lagerstätten of the vertebrae. Thus, the position of the Sclerorhynchoidei and newly found complete rhinobatid taxa from the Upper the Jurassic and Cretaceous guitarfishes (likeSpathobatis , Cretaceous of Lebanon allows the possibility of integral

*Author for correspondence: [email protected] basal group position. The phylogenetic relationsbasalbasalhips group group within position. position. batomorphs The The phylogenetic phylogeneticare yet even relations more relations hipships within within batomorphs batomorphs are are yet yet even even more more basal groupbasal position. groupcomplicated position.The phylogenetic when The fossilsphylogenetic relations are considered hipsrelations within becausehips complicatedbatomorphs withincomplicated their batomorphs fossil whenare when yet record fossils even fossilsare is yetmoreare mostly are evenconsidered considered restrictedmore because becauseto teeth, their their fossil fossil record record is mostlyis mostly restricted restricted to teeth,to teeth, complicatedcomplicated whendermal fossils when denticles are fossils considered and are occasionally, considered because theirbecause fin spinesfossil theirdermal recordanddermal fossil calcified denticles is recorddenticlesmostly vertebrae. andis restricted mostlyand occasionally, occasionally, Thus, restricted to teeth, the fin position to finspines teeth, spines of and the and calcified calcified vertebrae. vertebrae. Thus, Thus, the the position position of theof the dermal denticlesdermal Sclerorhynchoidei denticlesand occasionally, and occasionally, andfin spines the Jurassic fin and spines calcified and and Cretaceous calcifiedvertebrae.SclerorhynchoideiSclerorhynchoidei guitarfishesvertebrae. Thus, the Thus,and position(like and the the Spathobatisthe Jurassic ofposition Jurassic the and ,of andBelemnobatis Cretaceousthe Cretaceous guitarfishes guitarfishes (like (like Spathobatis Spathobatis, Belemnobatis, Belemnobatis SclerorhynchoideiSclerorhynchoideiand and several the Jurassic andfossils the Rhinobatoids) andJurassic Cretaceous and Cretaceous in guitarfishes the phylogeneticand guitarfishesand (likeseveral several Spathobatis tree fossils (like offossils batomorphs SpathobatisRhinobatoids) ,Rhinobatoids) Belemnobatis and, Belemnobatis inwithin thein the phylogenetic Rajiformes phylogenetic tree tree of batomorphsof batomorphs and and within within Rajiformes Rajiformes and severaland fossils severalis far Rhinobatoids) fossils from clear. Rhinobatoids) in the phylogenetic in the phylogenetic tree ofis batomorphs far istree farfrom of from batomorphs clear. andclear. within and Rajiformes within Rajiformes is far fromis clear.far from clear. In the Upper Cretaceous of Lebanon, the batomorphsIn theIn the Upper Upper are Cretaceous represented Cretaceous of by Lebanon,of 16 Lebanon, species the ofthe batomorphs batomorphs are are represented represented by by16 16species species of of In the UpperRajiformesIn the Cretaceous Upper belonging Cretaceous of Lebanon, to 9of genera Lebanon, the batomorphs and the4 families. batomorphsRajiformes areRajiformes representedThe Rajiformesare belonging representedbelonging by 16 phylogenyto species 9to bygenera 9 genera16 of species remains and and 4 families. of4 poorly families. The The Rajiformes Rajiformes phylogeny phylogeny remains remains poorly poorly RajiformesRajiformes belongingunderstood belonging to 9 genera with to the 9and genera “Rhinobatidae” 4 families. and 4 families.The forming Rajiformes Theunderstood aunderstood Rajiformespolyphyletic phylogeny with with phylogeny groupthe remains the “Rhinobatidae” (Brito“Rhinobatidae” poorlyremains and Dutheil, poorly forming forming 2004; a polyphyletic a polyphyletic group group (Brito (Brito and and Dutheil, Dutheil, 2004; 2004; understoodunderstood withBrito the “Rhinobatidae”with et al, the 2013; “Rhinobatidae” Claeson forming et al. aforming polyphyletic 2013) aand polyphyletic theBrito group Britoposition et (Brito al,et group al, 2013;of and2013;the (Brito ClaesonSclerorhynchoideiDutheil, Claeson and et2004;Dutheil, al.et al.2013) 2013) 2004; fluctuating and and the the position due position of theof the Sclerorhynchoidei Sclerorhynchoidei fluctuating fluctuating due due Brito et al,Brito 2013; etto al,Claeson the 2013; lack et Claesonof al. available 2013) et al. and well-preserved 2013) the position and the specimenofposition theto Sclerorhynchoideitheto of s.the lack Similarly,the lack Sclerorhynchoideiof availableof available the fluctuatingphylogenetic well-preserved well-preserved fluctuating due affinity specimen duespecimen of thes. Similarly,s. Similarly, the the phylogenetic phylogenetic affinity affinity of theof the to the lackto ofthe available lackLebanese of available well-preserved fossil Rajidaewell-preserved specimen and Cylobatidae specimens. Similarly, s.is LebaneseSimilarly,not theLebanese yet phylogenetic solved. fossilthe fossil phylogenetic The Rajidae affinityRajidae diversity and ofand affinity Cylobatidae ofthe Cylobatidaethe Lebaneseof the is notis not yet yet solved. solved. The The diversity diversity of theof the Lebanese Lebanese LebaneseLebanese fossil RajiformesRajidae fossil and Rajidae is Cylobatidae possibly and Cylobatidae underestimated, is not yet is solved. not compared yet TheRajiformes solved.Rajiformes diversity to The the is greatdiversityof possiblyis the possibly diversification Lebanese of underestimated, the underestimated, Lebanese event observedcompared compared in to theto the great great diversification diversification event event observed observed in in RajiformesRajiformes is possiblythe Upper is possiblyunderestimated, Cretaceous underestimated, (Guinot compared and compared to Cavin, the great 2015),tothe thediversificationthe Upper34 greatbyUpper lumping Cretaceous diversificationGeorges Cretaceous event the Kachacha majority(Guinot observed (Guinot event et andal.of observed in andLebanese Cavin, Cavin, in2015), 2015), by bylumping lumping the the majority majority of Lebaneseof Lebanese R & K the Upperthe Cretaceous Upperrhinobatids Cretaceous (Guinot in the and(Guinot genus Cavin, and Rhinobatos 2015), Cavin, by 2015),, lumpingwhereas byrhinobatids lumpingthetheirrhinobatids majority relationships thein the inmajorityof the Lebanesegenus genuswith ofRhinobatos the LebaneseRhinobatos modern, whereas species, whereas oftheir thistheir relationships relationships with with the the modern modern species species of thisof this rhinobatidsrhinobatids in the genus in the Rhinobatos genus Rhinobatos, whereas, theirwhereas relationships theirgenus relationshipsgenus with are are unclearthe unclear modernwith (Cappetta, the (Cappetta, species modern 2012; of species 2012; this Claeson Claeson of this et al,et al,2013; 2013; Brito Brito et al.,et al., 2013). 2013). genus are unclear (Cappetta, 2012; Claeson et al,anatomical 2013; Brito exploration et al., 2013).of the studied fossil organisms. 3. Results and discussions genus aregenus unclear are (Cappetta, unclear (Cappetta, 2012; Claeson 2012; etClaeson al, 2013; et al,Brito 2013; et al., Brito 2013). et al., 2013). Subsequently, complete redescriptions for these species The topology of this analysis is in general agreement with The outstanding preservation lagerstätten of the newly found complete rhinobatid taxa The outstanding preservation lagerstättenwere of theundertakenThe newly outstanding foundand will complete constitutepreservation rhinobatid together lagerstätten with taxa a full of thethe newly previous found analysis complete of Brito rhinobatid and Dutheil taxa (2004) and The outstandingfromThe outstandingthe preservation Upper Cretaceous preservation lagerstätten of lagerstättenLebanon of the newlyallows offrom the from foundthanalysis newlyethe possibility the Uppercomplete Upperfoundof their Cretaceous ofcomplete Cretaceous rhinobatidaffinities integral of rhinobatid theanatomical taxaLebanonof subjectLebanon taxaallowsof exploration allowsforecoming the th possibilitye possibilityClaeson of integralofet integralal., (2012), anatomical anatomical with a explorationbetter exploration resolution for the from the fromUpper the ofCretaceous Upperthe studied Cretaceous of fossilLebanon organisms.of Lebanonallows th Subsequently, allowse possibility the ofpossibility of theofcompletepublications. integralthe studied studied of redescriptionsanatomical integralfossil However, fossil organisms. anatomical organisms. weexploration fordiscuss these Subsequently, exploration Subsequently,herein species the attributionwere complete complete redescriptionsaffinities redescriptions of the for Lebanesefor these these species guitarfishes. species were were Nonetheless, it of the studiedof the fossil studiedundertaken organisms. fossil and organisms. Subsequently,will constitute Subsequently, completetogether completewith redescriptionsundertaken a undertakenfullof redescriptionsthree analysis shark-like forand andthese ofwill theirwill “Rhinobatids” constituteforspecies constitute affinitiesthese were species together species the together subject wereto withRhynchobatidae with ofa full a full analysis analysispresents of theirof markedtheir affinities affinities discrepancies the the subject subjectin the of relative of positions of undertakenundertaken and willforecoming constituteand will publications. constitute together withtogether However, a full with analysis we a fulldiscuss offorecominganalysis theirforecoming hereinand howaffinities of the their thepublications. attributionremainingpublications. affinitiesthe subject guitarfishes of However,the threeHowever,of subject shark-like are we of notwe discuss related discuss toherein otherherein the batomorphthe attribution attribution clades of threeof with three shark-likeother shark-like studies based on different forecomingforecoming publications.“Rhinobatids” publications. However, species However, we to discuss Rhynchobatidae we herein discuss the herein attributionand“Rhinobatids” “Rhinobatids”how therecent attributionth ofspeciese remainingthree species Rhinobatos. speciesshark-like of three toguitarfishes Rhynchobatidaeto shark-like Rhynchobatidae are not andrelated and how how to th e th remainingephylogenetic remaining guitarfishes methods.guitarfishes The are torpediniformsare not not related related to are to recovered “Rhinobatids”“Rhinobatids” species to species Rhynchobatidae to Rhynchobatidae and how andthe howremainingother thothere remainingrecent guitarfishes recent species species guitarfishes are Rhinobatos. notRhinobatos. related are not to related to as sister group to all batoids in the morphological analysis other recent species Rhinobatos. et al. other recentother species recent Rhinobatos. species Rhinobatos. 2. Material and method of McEachran and Aschliman (2004), Aschliman (2012b) and Claeson et al. (2012) as well as in the 2. MaterialLebanon, andwhich method was located in the North East part of 2. Material and method 2.Gondwana Material during and methodthe Late Cretaceous, is endowed with molecular analysis of Rocco et al. (2007) and Rocco 2. Material2. Material andLebanon, method and whichmethod was located in the North EastLebanon, Lebanon,partseveral of which GondwanaKonservat-Lagerstätten which was was located during located thein that thein Late thehave North Cretaceous,North yielded East East a part variety part is of Gondwanaof Gondwana(2013). On during theduring other the the Latehand, Late Cretaceous, Rajids Cretaceous, are considered is is the Lebanon,Lebanon, whichendowed was which located withwas in locatedseveral the North inKonservat-Lagerstätten the East North part Eastof Gondwana partendowed of endowed thatGondwanaof duringexcellently have with with yieldedthe severalduring preservedLate several a Cretaceous,theKonservat-Lagerstättenvariety Konservat-LagerstättenfossilsLate of Cretaceous, (Philip excellently is et al., is1993). that thatThe have have yieldedancestral yielded a groupvariety a variety of ofall excellentlyofbatoids excellently in the molecular study of Our ,( .(2012),( حاقل) .al حاقل)known), Lebanese AschlimanLebanese fossil etfossil al. localities (2012a) localities andare are NaylorHaqel Haqel et حاقلendowedendowed with severalpreserved with Konservat-Lagerstätten several fossils Konservat-Lagerstätten (Philip et al., that1993). have The that yielded best preservedhavepreserved knownbest a yielded variety known fossils Lebanese fossils aof Lebanesevariety excellently(Philip (Philip fossil of fossil et excellently al.,localitieset localities al., 1993). 1993). are areThe HaqelTheHaqel best best (known from ) from studies of ( علما علما morphological ساحل)the ساحل) the the), Cenom) the fromCenomanian, Cenomanian,- analysis and isand inSahel agreement Sahel Aalma Aalma with علماfrom حاقل() fromfrom ( (ساحل ),( نمورة) Haqel نمورةحاقل)) localities )fossil and )) and andand Nammoura localities areNammoura NammouraSahel Haqel Aalma areحجوال ) حجوال) best ) Lebanesefrom knownHjoula theHjoula Hjoula Lebanese Cenomanian,fossil نمورة)et (Philip al., ) 1993).and et Nammouraal., The 1993). best Theknownحجوال ) preservedpreserved fossilsHjoula (Philip fossils from 1980).) from the1980). Santonian Shirai (1996) and Brito and Dutheil (2004) as well as the علما( Cappetta,from (ساحل ) ,Cappetta علما );Aalma1887; 1887 ساحل) Cappetta,the ) fromCenomanian, the 1980). Cenomanian,the andthe Santoniananian, SantonianSahel and and Aalma Sahel(Davis, Sahel (Davis, Aalma نمورة) from ;1887( نمورة) ,SantonianNammoura ) and Nammoura (Davis حجوال)Hjoula ) and theحجوال) Hjoula the Santonianthe Santonian (Davis, 1887; (Davis, Cappetta, 1887; Cappetta, 1980). 1980). (Davis, 1887; Cappetta, 1980). molecular study of Pavan-Kumar et al. (2014) in considering The Phylogenetic affinities of 15 batomorph taxa the shark-like as sister group to all batoids. The Phylogenetic affinities of 15 batomorphfromThe thetaThexa PhylogeneticLebanese from Phylogenetic the outcrops Lebanese affinities affinities(including outcrops of 15 oftwo 15batomorph (includingnew batomorph species ta xa ta xafrom from Threethe the Lebanese shark-like Lebanese outcrops“Rhinobatids” outcrops (including (including from Lebanon, Gen nov. 1 grandis (Fig. 1a), Gen nov. 1 maronita (Fig. 1b) and The PhylogenetictwoThe newPhylogenetic speciesaffinities with affinitiesof 15exclusion batomorph of 15 of batomorph two taxa Sclerorhynchids fromtwo ta twothe xawithnew fromLebanesenew exclusion species species thespecies) Lebanese ofwithoutcrops twowith haveexclusion Sclerorhynchids exclusion outcrops (includingbeen performedof twoof(including species)two Sclerorhynchids Sclerorhynchids based have beenon species) species) have have been been performed performed based based on on performed based on type material and additional Gen nov. 2 tenuirostris are united together within a clade two new twospecies new with species exclusion with exclusion of two Sclerorhynchids of two Sclerorhynchids species)typetype material havespecies) material been and have performedand additional been additional performed based well-preserved well-preserved on based on and and complete complete material. material. The The latter latter material material was was type material and additional well-preserved andwell-preserved complete material. and complete The latter material. material The latter was material including Rhynchobatus, as found in Brito and Dutheil type materialtype andmaterialprovided additional and by additional “Memorywell-preserved well-preserved of Time” and completeprivate and collection complete material.providedprovidedwas whereas material.Theprovided by bylatter“Memory “Memory some byThe material “Memory latterspecimens of Time”of wasmaterial Time”of Time” privatewere private was private purchased collection collection collection or whereas whereas(2004) some some and specimens Claeson specimens et wereal., were (2012), purchased purchased with theor exclusionor of provided providedby “Memorydonated by “Memory of to Time” the Natural of private Time” History collection private Museum collection whereas of Denmarkdonatedsomewhereasdonatedwhereas specimens to someor thetosome to the Naturalthespecimens specimens wereNatural Lebanese purchasedHistory wereHistorywere University. purchased Museumpurchased orMuseum or ofA donated or setDenmarkof Denmarkof to65 the or toorPristis theto the Lebanese, Spathobatis Lebanese University. andUniversity. Sclerorhynchus A setA set of. 65ofThese 65 two new donated todonated the Naturalmorphological to the History Natural Museumcharacters History Museumof (among Denmark of which Denmark or to 11 themorphological are orLebanesemorphological Naturalnew) to the were HistoryLebanese University. charactersused charactersMuseum in University. a Acladistic (amongof set Denmark(among of 65 A phylogeneticwhich set whichor ofto 11 the65 11are Lebanese are studynew) new) were weregenera used used present in ain cladistic atwo cladistic out of phylogenetic the phylogenetic three characters study study identified by University. A set of 65 morphological characters (among Nishida (1990) (propterygium failing to reach the level of morphologicalmorphological charactersbased on characters 36(among taxa. Thewhich (among matrix 11 which are was new) 11compiled arewere new) used basedusing basedwere in on Mesquitea used cladisticon36 36taxa. in taxa. a3.10 cladisticphylogeneticThe The (Maddison matrix matrix phylogenetic was study wasand compiled compiledMaddison, study using using Mesquite Mesquite 3.10 3.10 (Maddison (Maddison and and Maddison, Maddison, which 11 are new) were used in a cladistic phylogenetic the nasal capsules; pectoral fin posterior corner not reaching based on based36 taxa. on The36 taxa. matrix The was matrix compiled was compiled using Mesquite using Mesquite 2010)3.10 (Maddison and 3.10 the parsimony(Maddison and Maddison, analysesand Maddison, were performed using TNT 1.5-beta (Goloboff et al., 2003). 2010) and the parsimony analyses were performed2010)study using andbased theTNT on parsimony36 1.5-beta taxa. The (Goloboffanalyses matrix was were et compiled al., performed 2003). using usingthe TNTlevel 1.5-betaof the pelvic (Goloboff fin) for et whichal., 2003). Rhynchobatidae 2010) and2010) the parsimony and the parsimony analyses analyseswere performed were performed using TNT using 1.5-beta TNT (Goloboff1.5-beta (Goloboff et al., 2003). et al., 2003). 3. Results3.Mesquite Results and 3.10 and discussions (Maddison discussions and Maddison, 2010) and the exhibit plesiomorphic state relative to the most other 3. Results and discussions parsimony analyses were performed using TNT 1.5-beta batoids. The third characters regarding the caudal fin TheThe topology topology of thisof this analysis analysis is inis generalin general ag reementagreement with with the the previous previous analysis analysis of Britoof Brito and and 3. Results3. andResults discussionsThe and topology discussions of this analysis is in general agreement(Goloboff with et theal., 2003).previous analysis of Brito and presenting well- differentiated dorsal and ventral lobes, is The topologyThe topologyofDutheil this analysis of (2004) this isanalysis andin general Claeson is in ag general etreement al., (2012), ag withreement withtheDutheil Dutheilpreviouswith a better the(2004) (2004) previousanalysisresolution and and Claeson ofanalysis Claesonfor Brito the et and ofaffinities al.,et Brito al., (2012), (2012), and of thewith with a better a better resolutiononly resolution marked for for the Genthe affinities affinitiesnov. 1 ofgrandis. theof the This character is Dutheil (2004)Dutheil andLebanese (2004) Claeson and guitarfishes. etClaeson al., (2012), et Nonetheless, al., with (2012), a better with it presents resolution a betterLebaneseLebanese marked resolution for guitarfishes.the discrepancies guitarfishes. affinities for the affinitiesNonetheless,of Nonetheless,inthe the relativeof the it presents it positions presents marked ofmarked discrepanciesmissing discrepancies in Gen in nov. thein the 2relative tenuirostris relative positions positions (holotype of oflacking the LebaneseLebanese guitarfishes. guitarfishes. Nonetheless, Nonetheless, it presents it markedpresents discrepancies marked discrepancies in the relative in the positions relative positions of of extremity of the tail) and not scored for Gen nov. 1 maronita that presents a slender caudal fin with a well developed dorsal lobe. Revision of the fossil batomorphs from the Cretaceous of Lebanon, Vol 3. No 2 July-December 2017 and their impact on our understanding of the early step 35 of the evolution of the clade

Figure 1. Habitus for guitarfishes from the Upper Cretaceous of Lebanon. (a) Gen nov. 1 grandis, well-preserved male in dorsal view from Haqel (LB-V-2015-01); (b) Gen nov. 1 maronita, well-preserved and complete male in dorsal view from Haqel (NHMD-74748); (c) Gen nov. 3 B nov., holotype in dorsal view from Hjoula (LB-V-2016-01).

Three additional characters (rostrum representing phies (antorbital cartilage directly jointing the inner rostral more than two third of the total neurocranium length; teeth most part of the propterygium; teeth with sharp transverse with well-developed median uvula; pectoral radials all crest that rises centrally into an obtuse and low angle or articulating with pro, meso and metapterygium) are cusp; posterior section of the propterygium extending supporting this clade in the present analysis together with behind the procondyle). Gen nov. 2 tenuirostris and Rhyn- two plesiomorphies (presence of horn-like anterior chobatus are recovered in one clade and sister to Gen nov. processes; presence of labial cartilages). Thus, the two new 1 based on two characters (rostral appendices covering the genera Gen nov. 1 and Gen nov. 2 are excluded from distal third of the rostrum; pectoral radials extending not Rhinobatidae family and assigned to Rhynchobatidae. Gen reaching the level of the nasal capsules) with the addition nov. 1 is recovered as sister group to Gen nov. 2 based on of one plesiomorphic character (radials of the mesop- two characters (rostral appendices covering at least half of terygium are made of three undivided segments and the the rostrum; pectoral radials extending as far as the level dichotomy is located at the half of the pectoral fins length). of the nasal capsules) with the addition of three plesiomor- Gen nov. 1 present two dorsal fins those unlike Rhyncho- 36 Georges Kachacha et al. R & K batidae are caudally located to the pelvic fin. In a result, University that had put all their facilities under our the Rhynchobatidae family definition needs to be disposal. This work was funded by the Carlsberg founda- reassessed. tion under the project 107057 Study of exceptional fossil The crown guitarfishes are recovered as sister group and the University of Copenhagen. to rajids in the 50% majority rule tree based on one synapomorphic character (occipital condyles wider than References the base of the synarcual lip) in addition to five characters Aschliman, N. C., Nishida, M., Miya, M., Inoue, J. G., (presence of labial cartilages; ribbed dorsal groundmass Rosana, K. M. and Naylor, G. J. 2012a. Body plan denticles; absence of alar and/or malar thorns; placoid convergence in the evolution of skates and rays scales uniformly present; suprascapulae articulated with (Chondrichthyes: ). Molecular Phylogenet- vertebral column) and one plesiomorphy (rostrum length ics and Evolution 63(1), 28–42. less than two third of the total neurocranium length). This Aschliman, N. C., Claeson, K. M. and McEachran, J. D. clade on the top of the tree including the crown guitar- 2012b. Phylogeny of batoidea. In: Carrier J, Musick fishes together with rajids is supported by one synapomor- J, Heithaus M (Eds), Biology of sharks and their phy (propterygium extending as far as the level of the relatives. Boca Raton, FL: CRC Press, pp. 57–94. nasal capsules) in addition to three plesiomorphies (ante- Brito, P. M., Leal, M. E. C. and Gallo, V. 2013. A new rior fontanelle tabular in shape and stops before reaching lower Cretaceous guitarfish (Chondrichthyes, Ba- the rostrum extremity or at the level of the rostral appen- toidea) from the Santana formation, Northeastern dices base; the antorbital cartilage is joining the inner Brazil. Boletim do Museu Nacional, Geologia 75, 1 margins of the propterygium; pectoral radials extending –13 far beyond the level of the nasal capsules). As a result, the Brito, P. M. and Dutheil, D. B. 2004. A preliminary sys- order Rhinopristiformes recently erected by Naylor et al., tematic analysis of Cretaceous guitarfishes from (2012) is not recovered in our analysis where the crown Lebanon. In: Arratia G. and Viohl G. (Eds), Meso- group of “Rhinobatidae” presents a closer relationship with zoic Fishes 3. Dr. Friedrich Pfeil, München, pp. 101 Rajidae than to Pristidae. The crown Lebanese guitar- –109. fishes Gen nov. 3 (Gen nov. 3 hakelensis, Gen nov. 3 Cappetta, H. 2012. Chondrichthyes. Mesozoic and Ceno- whitfieldiand Gen nov. 3 B nov. (Fig. 1c)) and Gen nov. 4 zoic Elasmobranchii: Teeth. Verlag Dr. Friedrich (Gen nov. 4 latus and Gen nov. 4 intermidius) are recovered Pfeil, München, pp. 512. in two resolved clades in a polytomy with Rhinobatos, Cappetta, H. 1980. Les sélaciens du Crétacé supérieur du Trygonorrhina and Zapteryx, where all are rooted by Liban. I: Requins. Palaeontographica Abteilung A, Iansan and Aptychotrema. Rhinobatos is excluded from 168, 69–148. the crown Lebanese guitarfishes in the strict consensus tree Carvalho, M. R. 1996. Higher-level elasmobranch phylog- and based on two characters (pectoral propterygium not eny, basal squaleans, and paraphyly. In: Stassney M. reaching the level of the nasal capsules; tail with stocky L. J., Parenti L. R. and Johnson G. D. (Eds), Inter- appearance and representing half of the total body length). relationships of Fishes. Academic Press, San Diego, Gen nov. 3 is supported by three characters (antorbital California, pp. 35–62. cartilage free from the propterygium; teeth with incipient Claeson, K. M., Underwood, C. J. and Ward, D. J. 2013. lateral uvulae; posterior section of propterygium extending Tingitanius tenuimandibulus, a new platyrhinid ba- behind procondyle) whereas Gen nov. 4 is supported by toid from the Turonian (Cretaceous) of Morocco and two characters (nasal capsules with a horn-like anterior the cretaceous radiation of the Platyrhinidae. Journal process; teeth with flat crown and lacking shark transverse of Vertebrate Paleontology 33(5), 1019–1036. crest). However three additional characters (pectoral radi- Compagno, L. V. V. 2005. Checklist of Living Chondrich- als extending as far as the level of the nasal capsules; thyan Fishes. In: Fowler, S. L., Cavanagh, R. D., proximal segment of the propterygium is rostrally to the Camhi, M., Burgess, G. H., Caillet, G. M., Fordham, jaws level; radials of the mesopterygium are made of three S. V., Simpfendorfer, C. A. and Musick, J. A. (Eds.), undivided segments and the dichotomy is located at the Sharks, Ray, and Chimaeras: The Status of the Chon- half of the pectoral fins length) provide ambiguous support drichthyan Fishes. Status Survey. International for a clade including Gen nov. 4 as sister-group to Rhino- Union for the Conversation of Nature, Gland, batos in the 50% majority rule tree. Therefore, the mono- Switzerland, pp. 401–423. phyly of Rhinobatidae is not recovered and the assignment Davis, J. W. 1887. The fossil fishes of the chalk of Mount of Gen nov. 3 and Gen nov. 4 to “Rhinobatidae” is consid- Lebanon, in Syria. Royal Dublin Society 3(2), 457 ered as doubtful. –636. Douady, C. J., Dosay, M., Shivji, M. S. and Stanhope, M. Acknowledgments J. 2003. Molecular phylogenetic evidence refuting We would like to thank Lionel Cavin, Gaël Clément and the hypothesis of Batoidea (rays and skates) as de- Emma Bernard respectively from the Natural History rived sharks. Molecular phylogenetics and evolution Museums of Geneva, Paris and London for their help in 26(2), 215–221. accessing to their type collections. We are indebted to the Fowler, S. L. 2005. Sharks, Rays and Chimaeras: The Natural History Museum of Denmark and the Lebanese Status of the Chondrichthyan Fishes. Status Survey. Revision of the fossil batomorphs from the Cretaceous of Lebanon, Vol 3. No 2 July-December 2017 and their impact on our understanding of the early step 37 of the evolution of the clade

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