Forey, LOsez-ARSARELLO, & MacLeno: New Lenore: FaoM Morocco 1 FIGURE 1. 1. Type locally of tLenidotes pankowskd sp. naw The Holotype ‘28 the Teour area. 2. Section through Kem Kem beds. The holotype and additional sicull ro may only be 2 ‘was nat found in situ and the locali Seekenes neat ania undaubledly cere fram the ferugiieus dandsiene herizan. Aber Forey and Grande (1998). Diagnosis. Lange Lepicotes reaching an estimated Jength of 4.6 m (based on body propertions of L. mantedi) in which the parietal is about half the length of the frontal; left pants larger than the Suiruecapulart on emha vide of eben orbit diam- ster small (12% of frontal-pariets! midline length): complex suture between the nasal process of the with the anterior infrsor- BMNH P.66856, partial head showing skull roof, cheek bones. preopercie, interopercie, part of the opercle and posttemporal, and partial lower jaw of left side. extrascapular series of right side, vomer and crushed braincase elements. Kem Kem Beds, southeastern Maroceo (see below, Locasty) Paratype. BMNH P.64126, skull roof and par chesk of left side, varner. Locality. Tafilalt Region, Kem Kern Basin, south Taouz (Figure 1). As is usual with specimens fn this region the precise locality and horizon unknown. Many specimens were found lying loc on the surface and detailed stratigraphic work h nat been done. Specimens from the Kem Ki Beds of Taouz srea, southeastem Morocco sherk fauna and is thought to represent a det dopesk ierene of al: 3868). Etymology. Species named after Mr Mi Pankowski of Rookie, Maryland. who kin donated the holotype specimen to The Natural HFIGURE 2.4 virtual model of Lepidotes pankowshi sp.nov. BMNH PG6858 (Holotype), This model was made using Panic emaen wads DIB inh esarier expos aa Seis tim sepa wantin i arse panies much morphological detail at possible, no atlempt has been made to fill smal holes in the scan of optimise the model meshes. 1. Virlual model with texturemapced digitel image of fie actsal specknen. This representation pro vides the best portrayal of the specimen indiuding smudges, stains and other imeguisriies toc ne to be represented at part Of the Scanned surface texture, 2. False-color mesh led model showing maximum tustace detail. Both fies areuriten in the ‘icf format, which can be read by the PC version of Micrasof’s Intemet Explorer. The sean is also available in DXF, STL and WAL formats enline which can alse be used to view the sean using third-party software. tory Museum, London. We are grateful tor his patronage of palasontology. Description. Material described here is preserved in perfect three dimensions. in onder to give the reader some idea of the rebustness and faninful- ness to life conditions, we give a rotatable laser image as Figure 2, produced by one of us (MIM. ‘The skull roofing bones conform in shape and pattern most closely to those in Lepicotes mantelll and 1. maximus. Some of the measurements are shown in Table 1. 48 pointed out by Jain (1983, p. 34) some of the measurements of skull roofing banes are difficult because of extreme asymmetry between ipsiiaieral pariners as well as the imegu- larity of suture lines. The midline suture length (froniala and perietals combined in F64126 (ihe lerger of he two specimens} is 235 mm. Assuming the propesions af the entire fish to be comparable with 2. mantel then this skull comes from a fish of about 1.6 m total kength. The holotype would nave been slightly smaller. ‘The fontals meet one another through ar almost straight mutual suture (Figure 3) as in L mantet! but unlike &. semiserratus where it it decidedly sinuous. The suture between frontal anc parietal is interdigitating and that with the dev. mopterolic i sinuous. The trontal is about twice at Jong as the larger of the panstais and the maximuurr width af the frontal is st the level of the dermosphe otic. Anieriorty the frontal is sutured ta the pre maxilla through a complex interdigitating suture such that the two bones ane firmly anchorec together. A band of tiny pores runs parallel anc close to the lateral margin of the frontal marking the path of the supracrbétal sensory canal. ‘The parietals are asymmetrical (s commor feature in Lepidotes species). In the two spect mens described here the lefl parietal is approxt mately 110 percent larger than the right in area. As dain (1983) noted in species of Lepidotes, fm le usual that in the majority of individuals of any one species either the right or left elernent is langes, bu Mat there may be in a few indivigusls showing TABLE 1. Same measurements of holotype and paratype of Lepicotes pankoushr sp. now. MS, midline suture tengit MPS, mutual frontal euture jength; FW, maximum frontal width; FL. maximum frental length: LPL maximum lengih bet navielal: RPL: maximum leig® right pecietal: OD, maximum diameter of arbi: FFL, ralien between the mazirrun frontal width and the macmum erttal length: PLIMPS, rate between the mamum parctal leg and the mutual for fal eudure length; OCUMS, ratio belween the maximum disrieter of orbit and the midline sulure length. Meaturerrertt are expressed in mm and ratios a8 percentages. as oMFS FW FL LPL RPL 00 FFL PUMFS ODS =e oo a Ta = = 2 coy ae Tze 235 10 = 17a as m a om 7 18%,FOREY, LOPEZARSARELLO, A MACLEOD! NEW LERIDOTES FROM MOROCCO FIGURE 3. Lapiivtes pankewski sp.nev. BMNH PS8856 [Holotype] skull i dorsal view wilh inlerpretive erawing Abbreviations: anit, anterior infraorbital; Ao, anterbital, Dpl, decmapteratic. Deph, decmasphenatic: Exec, extras eapuie: Lolf, foramen for olfactory nerve; Fr, frontal; Ne, nasal: Pa, parietal: Pra, premauilla: Pel, postiemporal, Se, supiaocbital, Sab, euborbital opposite size relationships. Of course, with only two individuals we cannot be certain if this species favours larpes left parietals. The dermopterotic reaches fonwand tp an end level with the maximum forward extent of the pan etal. Because the left paristal is larger than the Fight, mere is a compensation in the demmopterotcs sizes such that the nght is slightly wkier than the lef. Many tiny pores mun in. line close to the lateral edge of the dermopterotic marking the path of the ‘otic Sensory canal. ‘All of the roofing bones show an omament pattem of iow vermiculating ridges. Ganoine appears to be absent fom all skull bones. The absence of ganaine is unike ather species of Lent detes. Most Jurassic species of the genus have an extensive covering of ganeine on the skull bones. ‘Crelacecus epecies tend to have ganoine covering restricted to isolated tuberies on tne roofing and cheek bones. in the species described here even thet restricted ornament appears to be absent. We should add that westhering may hawe removed al races; Merefore we are cautious in cisiming that this species is totally devoid of ganoine armament There ere three extrascapulers on elther skde of the midline {only those of the right side are pre- seved in the holatype). The medisl and lsteralmost are both larger than the middle of the series, and in GMNH P.64126 tne medial Is larger than the tat- eralmost {in the holatype the medial is broken so the reiative size is unknown), The medial extras- capular is sutured to the parietal the isteraimost win the dexmopteratic: while me middie element sutures wath both the parietal and the dee mopterotic. in both specimens the dorsal tip of te post temporal can be seen to project beneath the mid- die member of the extrescapular senes and in MINH P6426 can be seen to reach the medial exirascapulsr. in BMNH P.64126 there are fragments of a median scale plus the first of the lateral scales. Al the anterior end of the skull parts of me pre maniliae can be seen in both specimens, but they are much better preserved in the holotype. Here they are extremely large snd stout (Figure 4). The anterior part of the premaxiis is broad and much thickened. ARhough the anterior edge is broken, the root canals for about 10 teeth in each premax-GURE 4. Lepidates pankewskii sp new. BMNH P.68856 [Holotype] skull in Jef lateral view with interpretive drawing. sbreviations: Ang, angular; ant.Int, antericr infacebital; Ac. arts osphenctic: Exec, extrascapul supercte; Op, opercis; Pa, pa woe. ; Den, dentary; Dpt, dermopterotic; Deph, dex af, foracnen far olfactory nerve; Fr, frontal, inf, ieltaorbital, lop, interepercie; Pap, ; Pena, premaxilia; Pll, posiiemporal; Se, supracrbital; Sob, suborbilal; Sop, subo-FOREY, LOPEZ-ARSARELLG, & MACLEOD: NEW LERDOTES FROM MOROCCO la can be seen. The nasal process Is broad and ong, and the dorsal surface af the posterior end of the process is tightly untied by an interdigitating suture to the underside of the frontal such that this joint is immoveable. An interdigitating suture is characterstic of Lepidotes (although this feature is aiso seen in other halosteans). The nasal process i perforated by the large olfactory foramen. The premaxilise of aither side remain separate: Remains of the night nasal can be seen in the holotype where it lies. in situ. itis small and kregu- lar in Shape although the trae shape is unknown because the anterolateral edge is broken. A line of pores crosses the nasal indicating the path of the Supracmia! canal. ‘The circumorbital end cheek bones are best preserved on the left side of the holotype (Figure 4). There is close agreement with the patier of me remaining bones seen in BMNH Pea126. The orbit is very saa compared with the Jengtn of the head, the maximum diameter being equal to about 12 percent mki-suture tength of the skull roof. it appears to be considerably larger in most omer species of Lepidetes examined here (eg. in L. semisevratus this ratio ks 30% while in 1 mantel it is 25%), except for i. raxo/ (see Gallo and Brito 2004, figure 3) and probably ¢ iatitrons (see recon struction of the skull in Jain and Robinson 1963, figure 1) ‘There are eight bones bordering the orbit. The two above the orbit are usually designated as supraomitais. a lange element at the posterodore.al position can be called the dermosphenotic: since it Cames the infraorbital sensory canal onto the root where presumably jons wit the otic sensory canal, and five further elements border the poste rior and anterior margins. of the orbit. The two cincu- morbitals lying behind the eye are eech equidimensionail; those beneath the aye are deeper than wide. The supraorbitals and the der- mosphenolic are sutured wh the skull roofing bones. The elements behind and beneath the eye are marked with & prominent ridge along the orbital margin. This ridge may suggest the path of tne main iateral line canal but it should be pointed out that there is & series of tiny pores scattered close to the distal margin of thes= bones that may mark the surface exits of the infraorbital sensory canal ‘Lying in front of the circumorbital series there are five more bones in the holotype, and these rep- resent the emiarior infraorbital and antorbital series (terminology of Lépez-Arbarello and Sferco in press}. In BMNH P64126 only the mest pasterior of these is preserved. There are four anterior infraorbite’s, wo of them Be against the late edge of the frontal such thal there is no g between the skull roof and the anterior infraorti series (cf, new semionatiform trom Germa Lépez-Arbarella and Sfereo in press}. The m antenor infracrbital is a small triangular bone wa the ameriommost is 8 curved element that | above the nasal process of the premaxilla and usually called the antorbital. Pores, presumal related to the infraorbital sensory canal, can geen in all of the infracmitats. They are most pro inent in the anteriormosi element as a line closely spaced pores lying near the lateral edge the antorbital Between the dermosphenotic and the cir morbital behind the eye there is 2 tiny diamor sheped bone thet must be considered es part the circumorbital series. ft is found in both spe mens where itis similany developed and must considered as a characteristic for the species. ‘There is @ series of eight suborbitals, seven which are sutured to the bones of tne infraort and antorital series. At least three ile in front the vertical level beneath the anteriar margin of t orbit. The three most anterior subosbitals sutur with the four anterior infraorbitals, which is unique feature among semionotforms. The m posterior of the suborbitals is sutured to the dern Sphenotic and the dermopteratic. A small sub bitai ties at the posteroventral extent of tne che Senes but it fais to reach the circumorbital series Parts of the opercular bones are preserved. the Jeft side of the holotype. The preoperele Prominent. M has an elongate vertical limb meet @ horizontal limb through an angle of about 121 The norzontal limd reaches forward to the level the jaw articulation, Scattered pores mark ine pi of the preopercular sensory canal. The interopercie is deep and appears. tigh sutured to the whole horizontal limb of me preop cle. Behind this tevel parts of the subopercie 3 operde be seen, and all appear clost sutured to one another. The ascending process the suboperdie is broad, as is the case in Lepidal maximus of L sews (contrary to L. mantel or ry semionatifarm from Germany, Lapez-Arbarello 2 ‘Sferca in press}. ‘A small part of the ieft lower jaw is preserv in tne nolotype but lite useful information can gleaned except that the jaw joint is located at tical level beneath the anterior margin of the a ‘The anguiaris deep and shaws 3 complex sinus dal suture with the dentary (ine antenor part which is missing. so it is not possible to see tdepth of the symphysis — 2 feature of same impor tance in distinguishing some Lepkiotes species irom others). Many small pores mark the path of the mandibular sensory canal running dose to the ventral edge of the jaw. COMPARISONS AND DISCUSSION The species described here is referred to the genus Lepidotes based an the following camibina- Hon of characters that can be seen in the spect mens: asymmetircal parietals: more than two anterior infracdbilals: a series of more than two suborbitals extending ventral to the omit; close overiap between the opercle, interopercie. subo- perce and propercie Although af these features are present in the type species Lepidotes efvensis, none of these characters alone is unique to Lepr- does species, and indeed the genus is in urgent seed of revision. There are well ower 150 nominal species described from deposits ranging from the Rhaetic through Cenomanian. Almost any large Mesozoic fish showing thick rhomboid scales. thick skull bones and robust grasping and crushing teeth runs the risk of being identified as a species of Lepidases. Many of these nominal species are based on very fragmentary materisl, paady construed and must be considered as nomina dubis. Others are differ- ent parts of the same species, and some have been associated (see Woodward 1895). Yet others have been given specific names simply because of their geographic or stratigraphic locations. Clearly this situation only emphasises the need for a com- prehensive review that must also include species refered to the genus Semionotus (another com- mon species-rich taxon often confused with Lept dates). An added dimension to this taxonomic uncertainty is the fact thet the lype species of the genus is Lepidotes elvensis (Blainwille 1818) from the Lower Jurassic of Germany, France and Eng- land. This form differs considerably trom many Upper Jurassic and Cretaceous species referred to the genus in the pattern of skull roof bones, num- bers of extrascapulars, cheek bones_ dentition and depth of mandibular symphysis (see Jain and Rob- inson 7963, Jain 1983). A is very likely therefore that many of the later species, including the taxon described here, will have to be referred to new genera. However, since this cannot be decided until much more revisionary work has been com- pleted we keep this taxon in the genus, preferring this action to creating additional names that may nat be justified. PALAEO-ELECTRONICA.C Despite the taxonomic unceriainty regard the limits of the genus the taxon descnbed fw resembles strongly some of the Upper Juras and Lower Grelaceous species that shaw stran whoral dentitions such as te Upper Jurassic £. i ws Agassiz 1837, 1. feombsi Jain and Robansi 1863, 1. maximus Wagner, 1863, 1 decoral Wagner, 1863, or Lower Cretaceous £. degemha Branco, 1885, 1. mantel Agassiz, 1833, and sourai Woodward, 1908. A tritorsl dentition was recagnised by J (1983) with the following characteristics: 1, co-os fled vomers: 2, tong tooth-bearing area on vom 3, corenoids thick with lange tooth-bearing areas; deep jaw symphysis; & inner teeth short (le_ | pedicitate) with iow convex or flat surtaces. Le dotes pankowskil shows characteristics 1, 2 ar (Figure 5). The jaw symphysis is unknown in i species. The caronoids are also unknown but it possible to speculate that, if found, they wo bear a large toothbearing ares because | opposing centinon upon the dermopalatine sho teeth arranged in at least sox bongituctinal rows. Among these ioral species the cheekbo series varies, Jain (1983) noted that within the £ wdotes species that he considered (usually the b ter known Jurassic and Cretaceous species) 5 a group containing species in whicn there are: 6 suborbitals (Jain 1983 named these bar cheskplates) arranged in = single row. This is contrast to 2 group containing species with 8 — suborbitals (= cheekpistes of Jain 1983). Lepiga Pankowskd falls into this latter group with at be eight suborbitats Jain further recognised that among those s% cies with the higher number of suborbitals sa had the bones sranged in a single row {e.9., manted) while others had & mosaic of bones (e. L souzal), rather similar to Piodetes nigenen Wenz, 1999. Lepldates pankowsAu complies 1 closely with those of the first group with the exc: tion that there is one suborbital wedged betwe the distal ends of two suborbitals. A similar patte ls shown by Lepidofes maximus (Jain 1985: pl 2. figures A, 6). Figure illustrates me cheek of pankewski alongside examples of two other trito Species recognised by Jain art Robinson (1963 In ef species studied so tar, the subarh sefes extends as far forward a6 the anterior le of the orbit. Lepiciotes pankowskl differs in that 1 senes extends well anterior to this level such 1 the anteriormost suborbiial reaches close to 1 ethmaid region. Furthermore. the most ante suborbitals in pankowski are peculiarly sutusForey, LOrSzARSARELLO, & Macleoo: New Lenoores From MoRDcCO midline FIGURE 58. Lepivotes pankowsii sp.nav. BMNH P-66856 [Holotyipe]. Ventral view of snout region ba shaw vamexi and demmnepalatine Geriitian. to the anterior infracrbitals. This feature is unique arnong ihe species of Lepijates and emang seri onotiforms in general. Another feature noted by Jain (1983) and Woodward (1895) is the increase in the numbers of extrascapuilars in inter occuring species. Most of Ine non-titoral species show a single pair of exirascapulars (eg., L semisemaius Agassiz, 1837. L deccanensis Sykes1851) while the tritoral species tend to show more (L. maximus — four pairs, L. mantatii — three or four pairs 1. souzai — three pairs). Lepidotes pankowskiY shows three pairs. A final observation nated by bom Woodward (1895) and Jain (1983) Is the fact that in later spe- cies the orbit decreases in relative size. The orbit of Lepiofes pankowshW ts parbcularly small and may result in the apparent increased prearbital lengm and, perhaps the hign number of antenor infreorbiais characterislic of this species. in sum i. penkowsil appears mast dosely similar to tritoral species such as L mantel and L. maximus in the dentition, disposition of the cheek pistes and the number of extrascapulars. it remains distinct in the high number of anterior infracrbtiais, the anterior extent and celationsh of the suborbitais, and the small size of the orbit The onder Semsonotiformes sensu Cisen z McCune (1991) is 2 monophyletic group inctud the lepisosteids, macrosemids and semionot The monophyly of the Lepisasteidae and the M rosemiidee are widely accepted, but ine semi otis most probably represent a non-monaphys sssemblage including all semsonotiforms that c nat be referred to one of the bvo monaptrytetic fe ies (see Lopez-Arbarello and Sferco (in press) @ brief historical overview). Nonetheless, recon of semionotid fishes in Atrica is patchy intriguing (Lopez-Arbaretc 2004, Lopez-Arban et al 2008). The oldest recerd in this continent is Sem otus capensis in the Early Jurassic Clarens Farr tion of South Africa (Woodward 1888). followed Lapidotes congolensis in the Middle Jurassic: teyville Beds of the Lualaba Senes in the Der atic Republic of Congo (Hussekof 1817, Sa ‘Seine 1955) and Lepidores tendagurwensis in Late Jurassic Upper Saurian Beds of Tencag (Arata and Schutze 1999). With the exceptior S. capensis. which is represented by numer relatively compiate and rather well-preserved spLi mantel FIGURE 6. Comparison af cheekbone pattems in three triteral species of Lepicotes. Lepidotes maximus based on dain (7885), L mantel based on Woodward (1316). See tent for ciecumsion imens, the other two species are only known from: Src ive Laks Nemec aagriaudelans omen in Ettéopia (Moody and Sutcliffe 1991, Goodwin et Lopez and Albian to Early Cenomanian Bokungo Beds inFOREY, LOPETARSARELLO, & MACLEOD: NEw LemnoTes FRO MORODCG the Democratic Republic of Cango, and the Hama ‘Koussou Basin of Cameroon: more detailed intor- mation in Murray 2000), but articulated remains are very rare. Apart from Lepidotes pankowskil mn. sp. described herein, compicte and well-preserved specimens previously identified in Lepidoes and currently recognized as 2 different taxon, Pliodetes nigeriensis, are known from the Aptian of Gadouts- ‘cua in Niger (Venz 1999). Additional complete and wallpreserved material identified as Lepiiotes manni of prooably Early Cretaceous age is reported from the Babouri-Figuil Basin in Camer- ‘con. but this species fs poorty understood and cur- rently under study (Giga Otero, personal commun. 2010). “yinerafore. she cody wail pmsarved maberikct semionotids is sparsely recorded trom the Early durassic ta the Cenomanian and represents quite different taxa. Even the species of Lepiotes. L. tendsguruensis and L pankowskil are probably not diosely related. Lepidotes fendagurvensis is. sirik- ingly similar to £. minor (ALA personal observa tions) from the Purbeck of England, which resembles Semionows im several features (McCune 1986). As previously discussed. pankowsid is most probably closely related to the large tfioral forms known from the Late Jurassic and Early Cretaceaus of central Europe. Lepidotes pankowski furthermore represents: the youngest confident recom of a semionotid giabally. Cther Canomanian or younger fossils identified in Lepidotes or simply referred to as ‘semionotids consist of isolated bones of, more te quenty scales or teatn. Almough at least some of these fossils might actually represent semionotids, many of them might tum cut to be lepisosteids stter thorough revision. ACKNOWLEDGMENTS We would lke to thank Regina Ellenbracht (formerly af the Palaeontology Conservation Unni, The Natural History Museum, London) for prepars- tion of both specimens described here. We would aise like to thank Phi Crabb (Photographic Unit, The Natural History Museum, London} for taking photographs of the specimens. This artide is 3 contribution ta Projecs DFG LO 1405/1-1 te 1-3. REFERENCES, dpesets, J $582, unierecchungen Ober te eaten Agattic, JLR, 1893 -1844, Recherches sur fet Pais sons Fassites, Petitpiesre, Neuchatel Aeambourg, ©. and Bertin, L 1956. On the fossil Sines found by Ms Gardiner in the Province of Ceara, in the North of Gran, Edinburgh Mew Philosophical Jou ist 30:82—84. Aeratis, G and Schultre, H.-P. 1959. Semionotiform fish from the Upper Jurassic of Tendaguru (Tanzania). Mitetungen sus dem Iduseam fir Neturkunde 20 Berlin, Geowissenschatiche Reibe, 2:136-153. Branco, WM. 1885. Ober eine neue Lepisotus - Amt aus Fe tiochversity. De. Friedrich Pfeil, Monctren. pes, ST-AB1~412. de Blaine, HL TB1@. Sur ket ichiyosies ou lee prois- sons fotsiles. Nouveau Dictomare | chistors Notmede, appiqude aux arts, 3 économie ruraie et bag myers repaid Forey. P-L. are Cavin, L. 2007. A new species af Clado- cycle (Teleosiei: Ichlhyedectifionnes) fom the Cenomanian of Moracen” Patseantoiogia Electron Ha TORI2ZA10p 1.1 MB htgcipalsecetecirore ica ong2007_ATSSVinde haem Forey, PL and Grande, L. 1998. An Alriean twin to the Brawilian Actinopterygii: Amide). cal Journal of the Linnean Society af Loncon, 123.178-185.