ORYCTOS,Vol. 1 :23-35,Octobre 1998 THEGENUS FURO (PISCES, HALECOMORPHI) FROMTHE UPPERJURASSIC PLATTENKALKE OF GERMANY Paul H. LAMBERS PaleontologischeWerkkamer, Biologisch Centrum RUG, Postbus 14,9750 AA Haren, the Netherlands. e-mail: phlambers@ biol.rug.nl Abstract : An overview of the speciesassigned to the genus Furo fowd in the German lithographic limestones of the Solnhofen-area(Bavaria) and Nusplingen (Baden-Wiirttemberg) is presentedand the monophyly of the Upper JurassicFuro is discussed.Six speciescan be recognized: 'F.' latim.anus,'F.' longiserratus, 'F.' microlepidotes, 'E' aldingeri, 'F.' angustus and'F.' miinsteri. Among these 'E' angustus and'F.' miinsteri form a monophyletic group, to which 'F.' aldingeri might be related as well. 'F.' longiserrarus might be closely related to the Ophiopsidae,whereas 'E' microlepidotes shows similarities with the Caturidae. The position of 'F.' latimanus remains to be determined. There are no indications of a monophyletic genusof Furo and the relationshipsof the Upper Jurassicfurids with the Lower Jurassicspecies of Furo remain to be examined. Key words: Eugnathus, Furo, Halecomorphi, phylogeny, Plattenknlke, Tithonian Le genreFuro (Pisces,Halecomorphi) du Jurassiquesupérieur d'Allemagne. Résumé : Les différentes espècesdu genre Furo enprovenancedes gisementsallemands à calcaireslithographiques des régions de Solnhofen (Bavière) et de Nusplingen (Bade-Wiirttemberg)sont présentéeset la monophylie du genre Furo du Jurassiquesupérieur est discutée.Six espècespeuvent être reconnues: '.8' Iatimanus, 'E longiserrattts, 'F.' microlepidotes,'F.'aldingeri, 'F.'angustuset 'F.'miinsteri.Parmi ces espèces,'F.' angustus et 'E'milnsteri consti- tuent un groupe monophylétique, auquel pourrait être rattaché 'E'aldingeri. 'F.'longiserratus se rapprocherait des Ophiopsidae et 'F.'microlepidotes montre des ressemblancesavec les Caturidae.La position de 'E' latimanus reste à être déterminée.Il n'existe aucuneindication sur la monophylie du genre Furo et les relations entre les furidés du Jurassiquesupérieur avec les espècesdu genre Furo du Jurassiqueinférieur doivent être étudiées. (traduit par la rédaction). Mots clés: Eugnathus,Furo, Halecomorphi, phylogénie, Plattenkalke, Tithonien INTRODUCTION England. The knowledge of the genus is mainly based on original descriptions from the 19th century The fossil fish genus Furo ( - Eugnathus,this and Woodwards Catalog of Fossil Fishes (1895a). F. name was preoccupiedby a genusof Coleoptera)is normandica, from the Toarcian of Normandy, des- known from the UpperTriassic of Lomb ardia(F. her- cribed by Wenz (1968), is the best known species. mesi, F. trottii. Alessandri,, 1910), severalspecies The monophyly of the genus has never been tested from the Lower Jurassic of England and France and it is therefore unknown whether all species assi- (Woodward, 1895a, b; Arambourg, 1935; Wenz, gned to Furo belong to this genus.The oldest known 1968)and the UpperJurassic of Cerin,France (Saint- representatives were originally described as belon- Seine,1949) and southernGermany. The type species ging to the genera Semionotus and Lepidotes and is F. orthostomus from the Lower Jurassic of referred to Furo by Allessandri (1910). 23 ORYCTOS,Vol. 1, 1998 I csgr q.n.d/^ Figure I a. Holotype of Furo longiserratus,specimen in the BayerischeStaatssammlung ftr Palâontologieund historischeGeologie, AS.Vil. I 136,Kelheim, Bavaria. b. Holotype of Furo latimanus,specimen in the BayerischeStaatssammlung ftir Palâontologieund historischeGeologie, AS.VII.262. Solnhofen.Bavaria. c. Holotype of Furo microlepidotes,specimen in the BayerischeStaatssammlung fùr Palâontologie und historischeGeologie, AS.V. I 1a,Eichstâtt, Bavaria. 24 LAMBERS- THEGENUS FURO In the Upper Jurassiclithographic limestonesof to group the Upper JurassicFuro into a monophyle- Bavaria and Nusplingen in Germany and Cerin, tic assemblagetogether with the Lower Jurassictype France,the following speciesof Furo havebeen des- species,all Upper Jurassicforms will be designated cribed (following the definition of Furo by 'Fu,ro'. Woodward, 1895 a): Furo aldingeri (Nusplingen; Heimberg, 1949), Furo vetteri (Bavaria?, Nusplingen; Heineke, 1906) Furo longimanus ?URO'LONGISERRATUS (B avaria),F uro longi se rratus (Bavaria,Nusplingen), Furo microlepidotus(Bavaria, Nusplingen) andFuro 'Furo' longiserratzs(Agassiz) (Figs. la,2) praelongus (Cerin, see Saint-Seine, 1949). 1843Pholidophorus longiserratus Agassiz, vol. II, Woodward (1895a) thought that Pholidophorus pt. I, p. 277, p1.38,fig. 2 (1843)(in 1833-1843) angustus, Pholidophorus elongatus and 1863Pholidophorus longiserratu,s, Wagner, p.664 Pholidophorus miinsterl belonged to Furo as well. 1863Pholidophorus brevivelis,rù/agner, p.664 Zittel (1887) placed Furo longisercatus,Furo longi- 1887Isopholis longiserratus,Zittel, p.216 manus and Furo miinsteri in a separate genus 1887Isopholis brevivelis,Zittel, p.216 Isopholis. He also included Pholidophorusbrevivelis 1895aEugnathus longiserratus (Agassiz), describedby Wagner(1863) in this genus. \iloodward,p. 301 ?19 14 Eugnathus lon gise rcatus (Agassiz),Eastman, The purposeof this paperis to give a short over- p. 410,pl. LXN, figs. 1, 2 view of the speciesof Furo from the GermanUpper 1949Eugnathus longiserratus Agassiz, Heimberg, Jurassiclocalities, to discusssome salient anatomical p. 97, pl. VIII, fig. 2 featuresand their implications on the phylogeny of 1966Eugnathus longiserratus (Agassiz), Schultze, Furo. A more detaileddescription of the speciesand fig. 19b an elaboratephylogenetic discussion will be publi- 1994Furo longimanus,Frickhinger, p.206,frg.434 shedin the future. As it appearsthat it is not possible - Figure 2 Skull of Furo longiserratus,lateral P+Dpt view. specimenin the Jura-Museum,SOS 3123. I Tithonianof Zandt.Bavaria. I sop Sor lo N lop Ao Mtp ec San F o Pm Ang Den mc G 25 ORYCTOS.Vol. 1. 1998 Holotype : Specimen AS.VII.I 136, Bayerische margin is embayed or only slightly notched, with a Staatssammlung fiir Palâontologie und historische small posterior process ventrally. The shape of the Geologie. Kelheim, Bavaria. maxilla is similar to that of primitive halecomorphs 'Fttro' longiserratus was originally described as a such as Brachyichthys (Woodward, 1895a), Conodus pholidophorid. This species was classified with the (pers. obs., a.o. specimen P.3645 in the NHM, poorly defined genus Isopholis by Zittel (1887). London), Heterolepidotes (pers. obs., a.o. specimen 'Furo' longiserratus is a slender, fusifoffn fish, P.2008in the NHM, London), Ionoscopus (pers. obs., with elongate head, pointed snout, high back, rather a.o. specimen 1903.I.64 in the Bayerische long dorsal fin placed posteriorly to halfway the body Staatssammlung fiir Palâontologie und historische length, narrow caudal peduncle, deeply forked caudal Geologie), Ophiopsis (Bartrâffi, I9l5), Teoichtlrys fin, the ventral rim forming almost a straight line (Applegate, 1988), Macrepistius (Schaeffer, 1960) from the anterior trunk to the posterior end of the and Lower Jurassic Furo (Woodward, 1895a; V/efz, lower lobe of the caudal fin and a scale-coverconsis- 1968).Most conspicuousis the presenceof a sensory ting of rhombic scales with serrated posterior mar- canal that runs through the maxilla. This is so far only gins, but in the throat region the scales are circular. known in the ophiopsids Ophiopsis and Teoichthys Most specimensmeasure between 10 and 15 cm stan- (Bartrâffi, I975; Applegate, 1988). The supramaxilla dard length, but the largest specimen that probably is half-oval. There areat least 14 branchiostegalrays. belongs to this specieshas a standardlength of 29 cm (specimen SOS.2I78a/b in the Jura-Museum, Eichstâtt). ,FURO'LATIMANUS The skull anatomy is of the general halecomorph type, with large first infraorbital, a single infraorbital 'Furo'latimanus (Agassiz) (Figs. lb, 3) behind the orbit, a broad, plate-like maxilla, with straight dentigerous border and which is expanded 1838-1843 Pholidophorus latimanus Agassiz, vol. dorsally in its posterior part with the dorsal margin II, pt I, 280(1843), pl. 43 (1838),(in 1833-1843) directed obliquely ventro-dorsally. The posterior 1863Pholidophorus latimanus, Wagner, p. 665 Figure 3 N Prn lol Skull of Furo latimanus, dorsalview, specimenin the Jura-Museum,SOS 2814. l.ryrI ..\. Tithonian t::::,, of Dollnstein. 'i! . '-.II:. Enp t-, \\ Bavaria. :r,!. \t I .:i'iJj . ' \l I Sm i I Enp I Ang \t So tl _- soc 'i O Sor Sy Pop /l Hyo r I ( ll 26 LAMBERS - THE GENUS FURO 1881 Pholidophorus latimanus, Vetter, p. 58 the short, weekly forked caudal fin are unique for this 1887 Pholidophorus latimanus, Zittel, p.216 form. 1895a Eugnathus latimanlts, Agassiz, Woodward, p. 302 ,FURO' 1966 Eugnathus latimanus (Agassiz), Schultze, fig. MICROLEPIDOTES 26 1994 Furo vetteri, Frickhinger, p. 206, fig. 435 'Furo'microlepidotes (Agassiz) (Figs. lc, 4,,5) Holotype: Specimen AS.VII .262, Bayerische 1833Uraeus microlepidotes, Agassiz, vol II, pt. I, p. Staatssammlung fiir Palâontologie und historische 12 (in 1833-1843) Geologie. Solnhofen, Bavaria. 1839-1843 Eugnathusmicrolepidotes, Agassiz, vol. IV p. 118 (1839),vol. II, pt. II, p. I04, (1843),(in 'Furo' latimanus was originally described as a 1833-1843) pholidophorid. It is a small (standard length up to 10 1863Eugnathus microlepidotes, Wagner, p. 673 cm) fish, with a round dorsal profile, a short skull 1895a Eugnathus microlepidotes, Agassiz, with a rounded or blunt snout, a triangular dorsal fin Woodward,p. 300 situated posteriorly to halfway the body-length, 1906Eugnathus