The Genus Furo (Pisces, Halecomorphi) from the Upper Jurassic Plattenkalke of Germany

Home , Calamopleurus, Caturidae, Caturus, Furo (fish), Halecomorphi, Ophiopsis

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 microlepidotes Ag., Heineke,p. 27, broad pectoral fin and a short, only weekly forked fig. 7, pL.V figs. 5-7; pl. VII, fig. 5 caudal fin. No specimens that clearly show the skull 1966 Eugnathusmicrolepidotes Agassiz, Schultze, anatomy in lateral view afe available. Nevertheless, figs.13 ,25 from examined specimens it appearsthat the maxilla 1994Eugnathus microlepidotes, Frickhinger, p. 433 is of similar shape as that of 'F'. longiserratus, but it Holotype: Specimen AS.V.1 lalb, Bayerische does not contain a sensory canal.The dorsal border of Staatssammlungfiir Palâontologieund historische the supramaxilla is slightly concave. The operculum Geologie.Eichst att, B avaria. is roughly quadrangular. There is one large supraor- Agassiz originally described this species as bital ventral and posterior to which many smaller Uraeus, a genus in which he also put speciesthat supraorbitals are present. The broad pectoral fin and were later referredto Pholidophoridaeand Caturidae.

Figure4 Skull of Furut rrtic'rclepiclote s, lateralview, II specimenin the tt Bayerische tl Staatssammlungftr il! Palâontologieund histori sche Geologie, t964.Xxrrr.554, Schernfeld, Bavaria.

1cm r RBr

27 ORYCTOS,Vol. 1, 1998

'Furo' microlepidotes is an elongate fish, of The triangular dorsal fin is placed posteriorly to rather large size (standardlength up to 35 cm), with halfway the body, the caudalfin is deeplyforked and an elongateskull with a blunt snoutin which the dor- symmetrical,very similar to the one of Caturus. sal and ventral border of the skull are more or less parallel. As in other halecomorphs there is one infraorbital posteriorto the orbit. Conspicuousin the skull is the slender,bar-like maxilla, which lacksthe 'FURO'ANGIISTIIS,'F(IRO'UÛWSfnnt dorsal expansionposteriorly but that is slightly dee- AND'FUR0'ALDINGERI pened downwardsinstead, with a clear notch in its posteriorborder and widely spaced,laterally com- 'Furo'angustus(Miinster) (Figs. 6a,b,c,7) pressedteeth. The shapeof the maxilla is very different from 1842Pholidophorus ongustus,Mtinster, p. 43 all other speciesof Furo and is in fact only known in 1848Pholidophorus Muensteri, Giebel, p. 208 the caturid genera Caturus and Amblysemius 1863Pholidophorus elongatus, \ilâgner, p.664 (Lambers, 1992, 1995).The supramaxillais sharply 1895aPhotidophorus elongatas, Woodward, p. 304 pointed anteriorly. The parietals are unequal in size 1895aPholidophorus angustu^ç, Woodward,p.477 with irregular,sinuous outlines. Parietals with irregular, sinuousoutlines are known in severalhaleco- Holotype: SpecimenAs.V[.1137,Bayerische morphs,but the big size differenceis only known in Staatssammlungfiir Palâontologieund historische Caturus (Lambers, 1992, 1994) and Osteorachis Geologie.Môrnsheim, Bavaria. (Gardiner,1960). The squamationconsists of narrow rhombic scales, at the ventral rim measurins not more than lmm.

Figure5 Skull of Fttrut Dpt rrtic'r'olepiclotes, // L ï, //ls St lateralview. il ,\l ! L. -scom skullroof in dorsalr'ieu'. /"{, PT {É 'qdrt . specimen it'ff/\ Jura-Museunt. ,:lt,-( -t ) È:r -4-\r/ ) È i /r) sos 305.1. -. Tithonian of Zandt, N Bavaria. R Ao

lo1 \ /--l

.RBr

28 LAMBERS - THE GENUS FURO ORYCTOS,Vol. 1, 1998

Mûnster (1842) described a small and slender the first branchiostegalrays which aredeeply seffa- fish with a long and slender pectoral fin as ted. The serrationsare directedpostero-ventrally in Pholidophorus angustus (Fig . 6a). Giebel (1848) the opercularbones. There are aboutnine branchios- briefly redescribed the species as Ph. miinsteri, as tegal rays. The supraorbitalsconsist of numerous Agassiz had already described a Ph. angustus from small and irregularshaped bones. the Jurassic of Poland. Wagner (1863) described a similar form as Ph. elongatus (Fig. 6b). He mentio- ' Furo' miinsteri(Agassiz) (Fig. 8) ned the similarity between these two species. The holotype of Ph. angustus Mtinster however, is an 1834Ophiopsis mtinsteri, Agassiz, p. 385 impression in the matrix only, lacking any trace of the 1843 Ophiopsismiinsteri, Agassiz, vol. II, pt. f, p. skull and the holotype of Ph. elongatus shows a 292,pt. II, p. 289(rn 1833-1843) disarticulated skull but has only part of the pectoral 1851 OphiopsisMiinsteri Ag., Wagner,p. 60 fin preserved. Probably for this reason he did not 1887Isopholis Miinsteri Ag ., Zittel, p . 216, ftg. 230 synonymize both species.The holotypes of the above 1895aOphiopsis miinsteri, Woodward, p. Il3 mentioned species,together with recently discovered material from other collections, allows a rather detai- Holotype : SpecimenAS.V[. 11 35, Bayerische led description of the species 'Furo' angustus.This Staatssammlungfiir Palâontologieund historische fish is an elongate, slender, small fish, that tapers Geologie. Kehlheim, B avaia. Figured is specimen gently towards the caudal fin, with a small skull with 1870.IV.2 (Bayerische Staatssammlung)from rounded snout. Conspicuous is the extremely long Wellenburg,near Kelheim,Bavaria, which hasbetter and slender pectoral fin and the short and slightly preservedskull and pectoralfin. bifurcated caudal fin which counts L9-20 rays only, in Agassiz ( 1834) erected a species,Ophiopsis contrast to the caudal fin of the other furids which miinsteri, based on the descriptionof scalesonly. containsabout 30 rays. Wagner( 1851) provideda more detaileddescription Characteristicsin the skull are the presence of of the holotype.Zittel (1887)referred the speciesto five supratemporalsand the posterior/ventral border the genusIsopholis and presenteda schematicdra- of the supratemporals,operculum, suboperculumand wing of the skull.

Figure7 Skull of Furut angustus, lateralview, the skull roof is seenin dorsal view, specimen in the collection ,''êC -R Kûmpel, Op Wuppertal. Pm Max

Cl J Ll ''1

L,' / /---'/--"'

--) r

5mm

30 LAMBERS - THE GENUS FURO

-il llllllll"tllli*i Figure 8 : Specimenof Furo m[insteri,specimenin the Bayerische Staatssammlungfiir Palâontologieund historischeGeologie, 1810.LY.2, Wellenburgbei Kelheim,Bavaria. Scrn

The species 'Furo' miinsteri is similar in shapeto limestone of Nusplingen, lacking the anterior part of 'F.' angustu* but with a standardlength of about 32 the skull. Additional material (PU.804401I2 from cm much larger in size. As in the latter it has a very Nusplingen, Baden V/iirttemberg, StaatlichesMuseum long and slender pectoral fin and a small caudal fin. fiir Naturkunde; fig. 10), displaying isolated skull In specimen 1870.IV.2 (Bayerische Staatssammlung bones allows a detailed description of the species. fiir Palâontologie und historische Geologie) seffa- 'Furo' aldingeri is a very slender and elongate tions on the posterior border of the operculum are fish, about 10 horizontal scale-rows broad, with visible. The vertebrae are well ossified ringcentra, conspicuously broad pectoral, dorsal, pelvic and anal unlike those of 'F.' angustus,which are composed of fins. The plate-like maxilla is widened dorsally, as in dorsal and ventral hemicentra. The caudal fin is assy- e.g. 'F.' longiserratus, but also has a rather rounded metrical in shape,âS the upper lobe is longer than the ventral deepening in its posterior third. The posterior lower lobe, whereas in 'F.' angustus both lobes are margin of the maxilla is deeply concave.The teeth on equal in size. The serrated posterior margin of the dentary and maxilla are slender and sharply pointed. posttemporals and the opercular bones in 'Fltro' The interoperculum has a posterior serrated margin, miinsteri and 'F.' angustus is unique among haleco- with more or less ventrally directed and widely spa- morphs. Some speciesof Ophiopsis have a denticula- ced serrations. There are several small and irregular ted posterior border of the supratemporals (Bartram, supraorbital. The parietals are rectangular and equal 1975). The long and slender pectoral fin and the in size. The small scalesbehind the skull are serrated small caudal fin are unique among halecomorphs as posteriorly, but halfway the body the scales lack ser- well. These characters unite 'F.' angustus and 'F.' rations, and the posterior margin is affow-shaped. miinsteri in a monophyletic genus. Five supratempo- The lateral line scales are deeply notched, showing rals is a derived condition, the number of supratem- two posterior projections. The extremely slender porals of 'F.' miinsteri is unknown. The assymetric body shape, and broad fins are unique for 'F.' aldin- shape of the caudal fin is unique for 'F.' miinsteri. geri. The shape of the maxilla and its posterior margin is different form the maxillae in other haleco- 'Furo' aldingeri (Heimberg) (Fig. 9a, b, c, 10) morphs (see above). The type of lateral line scales and the overall squamation is not seen in any other 1949 Eugnathus aldingeri, Heimberg, p. 95, pl. halecomorph. The lateral line scales ate usually not V[I, fig. 1, fig. 10 very conspicuous and the rhombic scales of haleco- 1966 Eugnathus aldingeriHeimberg, SchuItze,fig. 56 morphs usually have a serratedposterior margin. The serrated posterior margin of the interoperculum is Holotype: Specimen PV.I9495alb, Geologisches slightly different from the situation in 'F.' angustus, und Palâontologisches Institut of the university of as the serrations are less deep and the 'teeth' are more Tûbingen. Nusplingen, Baden-Wûrttemberg. widely spaced and not directed postero-ventrally. Heimberg described 'Furo' aldingeri after a single However, this difference might well be a preservatio- specimen in part and counterpart from the lithographic nal effect. 3r ORYCTOS,Vol. 1. 1998

Pcna

Figure 9 a/b. Holotype of Furo aldingeri, specimenin the Geologischesund Palâontologisches Institut of the universityof rûbingen, PY.19495alb,Nusplingen, Baden-wùrttemberg. c. Specimen of Furo aldingeri, specimenin the StaatlichesMuseum fùr Naturkunde Stuttgart, PU.8044Ol12, Nusplingen, Baden-Wiirttemberg.

32 LAMBERS - THE GENUS FURO

Figure 10.- Disarticulatedskull of Furo aldingeri,, from the specimenfigured in Fig. 9c; specimenin the StaatlichesMuseum ftr NaturkundeStuttgart, PU.804401 12, Nusplingen, Baden-Wûrttemberg. The skull has been rotated90' with respectto its original position.

--J--1 sol \l Scl Op Sop Sor fop 1cm

DISCUSSION big as between separategenera. Clear differences between all species are shown in overall body-form, Up till now 10 fossil fish speciesfrom the Upper skull shape, squamation, position, shape and size of Jurassic of Germany were assignedto the genus the fins. A few characters deserve mentioning. 'F.' Furo. On which groundsthe assignmentto this genus latimanus, 'F.' miinsteri and 'F.' aldingeri have large was basedis, however,unclear. The genusFuro is in two to three large supraorbitals, accompanied by fact a Liassic form and the knowledgeof the Liassic numerous small infraorbitals. Numerous small representativesis very incomplete.Apparently elon- supraorbitals are known form several halecomorphs, gate, fusifonn Jurassicfishes with rhombic scales e.g. Caturus (Lambers, 1992) and the amiiform with serratedmargin and rather heavy dentition were Calamopleurus (Maisey, 1991). This character is the- all placedwithin this genus.In publicationsauthors refore no usefull indicator of phylogenetic relation- often refer to the genusFuro (e.g. Schultze 1966, ship. 'F.' Iongiserratus has a sensory canal running 1996; Lambers, 1995) without taking into account through the maxilla, a condition that is only known in that the genus,âs presentlyknown, is not monophy- Ophiopsidae. However, this species lacks the deep letic, probablya polyphyleticassemblage and as such infraorbitals and the sensory line continuing into the has no phylogeneticvalue. From my brief overview caudal fin, known in Ophiopsidae (Bartrâffi, L975), of the Upper Jurassicforms from Germanythe ana- nor does the maxilla have a posteriorly directed ven- tomical diversity of the speciesis apparentand is as tral hook anterior on the maxilla, as known in 33 ORYCTOS,Vol. 1, 1998

Ophiopsis (pers. observ. on several specimens,e.g. Mr. Kûmpel (Wuppertal) and Mr. H. Tischlinger 1938.58in the GeologicalMuseum, Copenhagen, see (Stammhamm)for making their private collections also reconstructionsin Bartram, 1975). 'E' longima- availablefor study.I thank dr. G. Viohl (Eichstâtt)for nus is, in skull anatomy, similar to the general primi- his hospitality to provide me the facilities to study tive halecomorphpattern, but in body-form and shape fossil fishes in the Jura-Museum.This researchhas of the fins the speciesis different from all other spe- been financed by the Alexander von Humboldt- cies. 'E' microlepidoteshas a bar-like maxilla that is Stiftung (Bonn). similar to the one in Caturidae and the irregular parietals are as in Caturidaerather than in other furid spe- ABBREVIATIONS cies.The very narow scalesmake this specieseasily Ang: Angular distinguishablefrom the other species.Its vertebral Ao: Antorbital column remainsunknown, it is not known whetherit ba: branchial arches has the broad haemal spines typical for caturids Cl: Cleithrum (Lambers,1995).'F.' angustusand'E' mùnsterishare Cor: Coronoid an unusuallylong and slenderpectoral fin, a relative- Den: Dentary ly small and blunt-snoutedhead, serratedposterior Dpt: Dermopterotic margins of suboperculum,interoperculum and bran- ec: ethmoid commissure chiostegalrays and a small caudalfin. The five supra- Ecp: Ectopterygoid temporalsof 'F.' angustusare unique amonghaleco- Enp: Entopterygoid morphs.The supratemporalsof 'F.'miinsteri are unk- F: Frontal nown. The upperlobe of the caudalfin of this form is G: Gular much longer than in any other Furo.T\ese two spe- Hyo: Hyomandibular cies probably belong to the samegenus . 'E' aldinge- lollolfIo2: Infraorbital ri is easily distinguishableby its extremely slender Iop: Interoperculum body-shape,very large pectoral, dorsal, pelvic and LF,t Lateral ethmoid anal fins, the shapeof the scalesand the lateral line Max: Maxilla scales,and the shapeof the maxilla with its very deep mc: mandibularcanal notch and round downward expansion posteriorly. mp: mandibularpit-line The serratedposterior margin of the suboperculum Mtp: Metapterygoid might indicate a relationship with 'E' angustus/'E' N: Nasal miinsteri. Apparently the Upper Jurassicspecies of Op: Operculum 'Fltro'do not belong to a monophyleticassemblage. P: Parietal They represent several distinct genera. It is not Par: Prearticular known whether any of the Upper Jurassic 'Furo' Pm: Premaxilla belongsto genusFuro as charccteizedby the Lower Pop: Preoperculum Jurassictype species.Their relationshipto the Lower Psp: Parasphenoid JurassicFuro rcmainsto be determined. Pt: Posttemporal Q: Quadrate R: Rostral ACKNOWLEDGEMENTS Rar: Retroarticular RBr: Branchiostegalrays I thank Dr. R. Boettcher (Naturkunde-Museum, San: Surangular Stuttgart),R. Schoch(Tiibingen), Dr. G. Viohl (Jura- sc: sensorycanal Museum, Eichstâtt), Dr. P. Wellnhofer (Bayerische Scl: Supracleithrum Staatssammlung fiir Historische Geologie und Scl: Scleroticring Palâontologie)and Dr. J. de Vos and J. van Veen scom: supratemporalcommissure (TeylersMuseum, Haarlem) for giving me the oppur- Sm: Supramaxilla tunity to study specimensunder their care. I thank So: Supraorbital 34 LAMBERS - THE GENUS FURO

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