This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions CLEMENT and BOISVERT-DEVONIAN LUNGFISH FROM BELGIUM 277 tra. In addition to his incorrect taxonomic attribution, Lohest idae Berg, 1940 (including Fleurantia and Jarvikia); and Rhyn- misinterpreted the operculum as a scapula, the cleithrum as a chodipteridae Moy-Thomas, 1939 (including Rhynchodipterus, coracoid, and the E bone as an isolated rib (Fig. 2A, B). How- Griphognathus, and Soederberghia). Schultze (1993) defined the ever, he accurately identified a pleural rib (Fig. 2A, B). Rhynchodipteridae as including at least Soederberghia, Jarvikia, and Fleurantia. Later, Schultze (2001) presented a cladogram of SYSTEMATIC PALEONTOLOGY Devonian dipnoans that included a radiation of denticulated forms: Barwickia [Fleurantia + Rhynchodipteridae], in which included SARCOPTERYGII Romer, 1955 Rhynchodipteridae Griphognathus [Rhynchodipterus + The and affinities of the DIPNOMORPHA Ahlberg, 1991 [Soederberghia Jarvikia]]. monophyly DIPNOI 1845 Rhynchodipteridae have been reviewed by Ahlberg et al. (2001), Muiller, who that be unrelated RHYNCHODIPTERIDAE Moy-Thomas, 1939 tentatively suggested Griphognathus may to Rhynchodipterus and Soederberghia, but regarded Rhyncho- Remarks-Campbell and Barwick (1990) proposed that the dipterus and Soederberghia as most closely related to each other. denticulated lungfish lineage should be recognized as suborder However, Friedman (2003b) considered this suggestion prema- Uranolophina which incorporates four families: Uranolophidae ture and suggested that the Rhynchodipteridae, if defined as Miles, 1977; Holodontidae Gorizdro-Kulczycka, 1950; Fleuranti- including only Soederberghia, Rhynchodipterus, and Griphogna-

FIGURE 2. Soederberghiasp. indet. Modave, Liege Province, Belgium, upper Famennian,Upper Devonian. Liege University, paleontology collection no. 5390a,b. A, no. 5390a,operculum, parts of the shouldergirdle and axial skeleton of a same individual;B, no. 5390b, counterpartof the same specimen.Abbreviations: CI, right cleithrum;Eb, left E bone; i.c, internalcrest of the cleithrum;?ns, ?neuralspine; Op, left operculum; pbl, postbranchiallamina of the cleithrum;pr, pleural rib; ?Ano, ?anocleithrum;v, vertebra;vl, ventral lamina of the cleithrum.Grey shading representspreserved bones. Scale bars equal 10 mm.

This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions 278 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 26, NO. 2, 2006 thus, may be paraphyletic with respect to Fleurantia and Jarvikia. ?Neural Spine-A long, straight, and convex spine-shaped Numerous derived characters suggest that Soederberghia is more bone (Fig. 2A, B) is tentatively interpreted here as a neural closely related to Fleurantia and Jarvikia than it is to either spine. Rhynchodipterus or any of the species of Griphognathus (Fried- Vertebrae-Eight vertebral centra are preserved in specimen man, 2003a). Phylogenetic and taxonomic questions concerning no. 5390a, b (Fig. 2A, B). For some of them, the imprint of their long-snouted lungfishes are still pending, so here we accept the concave anterior and posterior faces shows a small pit corre- content of Rhynchodipteridae as defined by Miles (1977): Soeder- sponding to the notochordal canal (Fig. 2A). Despite their rela- berghia, Griphognathus, and Rhynchodipterus. tively good preservation, no ventro-median or dorso-median grooves and canal-like pits, known to be present in Griphogna- thus and and Jarvikia (Jarvik, 1952), cf. SOEDERBERGHIA 1959 (Campbell Barwick, 2002) Lehman, can be observed. The isolated vertebrae housed in the Louvain- (Figs. 1, 2A, B) la-Neuve Catholic University (specimens no. PVL 10.634, 10.635, Type Species-Soederberghia groenlandica Lehman, 1959. 10.636, and 10.637), although larger, display the same morphol- Age-Upper Famennian, Remigolepis series; East Greenland. ogy as those of the specimen no. 5390a, b. Diagnosis-See Ahlberg et al., 2001. Referred Specimens-Libge University paleontology collec- Discussion tion no. 5390a, b, remains of one individual (cleithrum, opercu- as evi- lum, E bone, pleural rib, vertebrae, ?neural spine, ?anoclei- The E bones of this lungfish were undoubtedly paired left thrum). Louvain-la-Neuve Catholic University collection no. denced by the fact that the E bone, considered here as the PVL 10.634, PVL 10.635, PVL 10.636, PVL 10.637, remains of one, is asymmetrical and is clearly not a median element. Fur- isolated vertebrae. thermore, the straight margin of the isolated left E bone indi- The Locality and Horizon-Modave, Liege Province, Belgium; cates that it was medially in contact with the right E bone. in upper Famennian, Upper Devonian. presence of a single median E bone is known Jarvikia (Lehman, 1959; Campbell and Barwick, 1990; Friedman, 2003a) Morphological Description and Andreyevichthys (Krupina, 1987) but the fusion between the two bones seems to be highly variable since specimens of the or Specimen no. 5390a, b consists of disarticulated elements of Frasnian species Fleurantia denticulata display either unique the skull roof and of the pectoral girdle, as well as vertebrae and paired E bones (Graham-Smith and Westoll, 1937; Cloutier, other bones of the axial skeleton. Dermal bones are thin and 1996). The general shape of the E bone in the Belgian specimen of devoid of cosmine. Considering that fossils are rare in this local- is very similar to that of Soederberghia groenlandica ('Ptr' ity and that all elements in the block belong to a small lungfish, Lehman, 1959:figs. 2, 4, 14, pls. 2-4, 5A; Campbell and Bell, we assume that they all belong to the same individual. 1982:fig. 3) and Soederberghia simpsoni (Ahlberg et al., 2001:figs. all Left E Bone-Part of this elongate bone (Fig. 2A, B, Eb) is 2A, B, 3A). In Griphognathus sculpta (Schultze, 1969:fig. 16), missing but its imprint allows a complete reconstruction of its bones of the skull roof, including the E bone, are much smaller in shape. The surface of the bone is gently convex and its external than the operculum. This contrasts with the condition found the surface, which shows no sensory canal, is ornamented with elon- the specimen from Belgium, where the E bone is longer than gated ridges whereas its internal surface is smooth. One of its operculum, as in Rhynchodipterus elginensis (Schultze, 1969) and long margins is straight, while the other is embayed by three Griphognathus whitei (Miles, 1977). The operculum is very simi- notches. The straight margin is most probably the medial one, lar to those of Griphognathus sculpta (Schultze, 1969:pl. 4:1), and would have articulated with the right E bone in life. The Soederberghia simpsoni (Ahlberg et al., 2001:figs. 2A, B, 3A) and and embayments in the lateral margin would have accommodated S. groenlandica (Lehman, 1959:fig. 18, pl. 21A). Its elongate bones bearing the supraorbital sensory canal. The posterior mar- rectangular shape differs from the rounded operculum of Rhyn- gin of the E bone is short and rounded, whereas its anterior chodipterus (Ahlberg et al., 2001). border is flared and bifid. The dorsal lamina of the cleithrum is elongated and rectangu- as in Left Operculum-This bone (Fig. 2A, B) is very thin and lar and the ventral lamina is short and medially directed unornamented except along its anterior margin, which bears Griphognathus sculpta (Schultze, 1969:fig. 18) and G. minutens some tubercles and two long ridges. The operculum has a rect- (Schultze, 1969:figs. 19, 23). The shoulder girdle of Soeder- Jar- angular shape, antero-posteriorly elongated, with a rounded pos- berghia is poorly known (Ahlberg et al., 2001) and that of terior margin. vikia is unknown. Right Cleithrum-The dorsal lamina of the right cleithrum Only four dipnoan genera are known to possess ossified disc (Fig. 2A, B) is elongate and rectangular in shape. The impression centra, a derived feature among lungfishes (Ahlberg and Trewin, of the external side suggests the presence of fine, vermiform 1995), and all are Late Devonian in age. Soederberghia is known ornament. A strong internal crest (Fig. 2A) is present along the from the upper Frasnian or lower Famennian of Australia dorsal portion of its anterior margin. This crest slightly bends in (Young, 1999; Campbell and Bell, 1982; Ahlberg et al., 2001) and a posterior direction to end in the central region of the base of the upper Famennian of Greenland and Pennsylvania (Jarvik, the dorsal lamina. The region anterior to this crest is most prob- 1952; Lehman, 1955, 1959; Schultze, 1970; Ahlberg et al., 2001). ably the internal side of a well-developed postbranchial lamina Jarvikia is found in association with Soederberghia in the upper (Fig. 2A). The ventral lamina (Fig. 2A, B) is medially directed Famennian of Greenland (Jarvik, 1952; Lehman, 1955, 1959; and appears in section. It is half the length of the dorsal lamina. Schultze, 1970). Griphognathus is known from Frasnian marine No trace of scapulocoracoid is visible. deposits of Australia (Pridmore and Barwick, 1993; Campbell ?Anocleithrum-A rounded, rectangular bone (Fig. 2A) and Barwick, 1988, 2002), Latvia, and Germany (Gross, 1956; shows a thick and slightly sinusoid margin. It does not appear to Schultze, 1969, 1970; Miles, 1977; Rosen et al., 1981). Finally, be a scale and its shape is similar to the anocleithrum of Gri- Adelargo is known from the upper Famennian of Australia (Jo- phognathus sculpta (Schultze, 1969:fig. 13, pl. 4:2). hanson and Ritchie, 2000). The genus Rhynchodipterus is con- Pleural Rib-A long and curved element (Fig. 2A-B, pr) is sidered as having possessed poorly ossified centra that were con- present beneath the operculum. Its proximal extremity is en- fined to the anterior end of the column (Ahlberg et al., 2001). larged and flattened, consistent with the morphology of a pleural Renewed study of Rhynchodipterus suggests it has disc centra, rib. their posterior extension being unclear on the unique specimen

This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions CLEMENT and BOISVERT-DEVONIAN LUNGFISH FROM BELGIUM 279

(Friedman, 2003a). Only three centra are preserved in Soeder- Morphological Description berghia simpsoni (Ahlberg et al., 2001:figs. 2C, 3B). These ante- riormost elements of the vertebral column of this species are The shape of the lower jaw is very distinctive. It is very elon- nevertheless considered by Ahlberg et al. (2001) as disc centra gated with a long and well-developed dermal retroarticular pro- rather than incomplete arc centra. The vertebrae of Adelargo are cess (Fig. 3), a well-marked rounded depression for the glenoid well preserved, with notochordal canals of variable width (Jo- (see Fig. 3), a hook-shaped dermal ascending process (see Fig. 3), hanson and Ritchie, 2000:fig. 3D, E), but the notochordal canal and a very long and straight mandibular ramus. The anterior- is never completely occluded as in the Belgian specimen (Fig. most part of the lower jaw, including the symphysis, is missing. 2A, B). The vertebrae of the latter are very similar to those of Nevertheless it is obvious that the elongated part of the lower Soederberghia (Jarvik, 1952:pl. 8; Lehman, 1959). Although jaw was composed predominantly of the mandibular ramus (as in longer and more massive than the vertebrae of those of Soeder- Griphognathus, Fleurantia, and Soederberghia groenlandica). berghia from East Greenland (Jarvik, 1952:fig. 16A, B; Lehman, This differs from the condition in Rhinodipterus, in which elon- 1959), the vertebrae of the Belgian specimen are not longer than gation of the lower jaw is achieved through lengthening of the high like some vertebrae of Jarvikia (Jarvik, 1952:pl. 8:3, 5; symphysis (Schultze, 1992). Lehman, 1959). Jarvikia arctica has very distinctive spool-shaped The lateral side of the left lower jaw shows many similarities centra that differ from the more disc-shaped ossifications of with Soederberghia groenlandica (Lehman, 1959; Schultze, 1969; Soederberghia and Griphognathus. It is not known whether the Friedman, 2003a). The large notch on the top of the dermal other nominal species of Jarvikia, J. lebedevi (Krupina, 1999), ascending process (Fig. 3) is rounded and well marked as in possessed ossified centra (Friedman, 2003a). Dipterus valenciennesi (Jarvik, 1967), Griphognathus minutidens Despite its incompleteness, the rhynchodipterid specimen (Schultze, 1969:pl. 6:5, fig. 3) and Soederberghia groenlandica from Belgium-Lohest's 'amphibian'-is considered here to be- (Friedman, 2003a:fig. 16A). According to different authors, this long to Soederberghia. Among all dipnoans with stout disc-like notch accommodated a branch of the trigeminal V nerve centra, this specimen differs from the late Famennian Jarvikia by (Lehman, 1959); both branches of the trigeminal V and the facial the presence of separated E bones and antero-posteriorly com- VII nerves (Schultze, 1969; Pridmore et al., 1994); or only a pressed centra, from the late Famennian Adelargo by very thin branch of the facial VII nerve, with the trigeminal V entering the dermal bones and ring-centra vertebrae without well-marked no- lower jaw more anteriorly between the prearticular and external tochordal openings, and from the Frasnian Griphognathus by an dermal series (Miles, 1977; Jarvik, 1967, 1980; Friedman, 2003a). E bone longer than the operculum. Oral and mandibular sensory canals are enclosed within the Compared to the very large Soederberghia groenlandica from bones but their posterior openings are obvious. Posterior fo- Greenland (Lehman, 1959) and from the Cloghnan Shale, Aus- ramina for these sensory canals are situated close to one another. tralia (Campbell and Bell, 1982; Ahlberg et al., 2001), the Bel- The posterior opening of the oral canal (Fig. 3) is located near gian rhynchodipterid is very small. However, it is similar in size the dorsal margin of the lower jaw below the posterior end of the to S. simpsoni from Canowindra, Australia and S. groenlandica glenoid fossa. The posterior opening of the mandibular canal from the Catskill Formation, Pennsylvania (Ahlberg et al., 2001). (Fig. 3) is situated just below, far from the ventral margin of the The two species of Soederberghia, i.e., S. simpsoni from Aus- lower jaw. A deep groove, enlarging posteriorly, is present pos- tralia and S. groenlandica from Greenland and Pennsylvania terior to the opening for the oral canal. This groove also accom- (and seemingly from Australia) are defined on the basis of dif- modated the oral sensory canal. The disposition of these poste- ferences in the pattern of dermal bones of the skull table (Ahl- rior openings of the sensory canals is very similar to that of berg et al., 2001). It is therefore impossible to propose a generic Griphognathus minutidens (Gross, 1956:fig. 25B, C, H; Schultze, assignment for the Belgian specimen. We attribute it to a rhyn- 1969:fig. 3). The opening for the mandibular canal is more an- chodipterid cf. Soederberghia. teriorly situated, as well as nearer to the ventral margin of the infradentaries, in Griphognathus sculpta and G. whitei (Schultze, A RECENT THE 1969:figs. 13A, 16C; Miles, 1977:fig. 99; Campbell and Barwick, DISCOVERY CONFIRMS PRESENCE The OF SOEDERBERGHIA 1987:fig. 14B). mandibles of Soederberghia groenlandica fig- ured by Lehman (1959:fig. 19; pl. 9A), Schultze (1969:fig. 39), and Friedman (2003a:fig. 17A, B) show a sensory canal (man- With the aim of more and associated verte- finding tetrapod dibular or oral canal) running far posteriorly to the glenoid fossa. brate remains, recent field in the Famennian of Bel- collecting This may represent a specific character of S. groenlandica, or has resulted the of the Strud which gium in rediscovery quarry, may instead represent individual variation of the kind seen in had been abandoned for over a This the century. quarry yielded Griphognathus minutidens (Gross, 1956:figs. 25B, C, H). et misidentified ichthyostegid jaw fragment (Cl6ment al., 2004) A thick but relatively short crest is present along the ventral as a fish Lohest and the horizon that by (1888a), probable pro- margin of the lower jaw below and slightly anterior to the dermal duced this was rediscovered in specimen October 2004. Precise ascending process. The dorsal border of this large crest forms of this dating locality by stratigraphical correlation and miospore the insertion area for the adductor mandibulae muscles (Fig. 3; biostratigraphy indicates a late middle Famennian age for this Schultze, 1992:fig. 13A). The same longitudinal crest is present in Thorez and locality (Jacques Maurice Streel, pers. comm., Janu- Griphognathus but it is more strongly developed (Miles, 1977:fig. New ary, 2005). material found in this tetrapod-bearing stratum 95C-M). The posterior part of the lateral side of the retroarticu- includes Holoptychius scales, dipnoan dental plates, indetermi- lar process also presents a broad and rounded ventral region. nate bone fragments, and an elongated mandibular ramus, which The dorsal border of this thickening may have formed the inser- we refer to the genus Soederberghia. tion area for a retroarticular/hyomandibular ligament (Fig. 3) which is known to run between the lateral face of the hyoman- dibular and the lateral face of the retroarticular in coelacanths SOEDERBERGHIA cf. S. GROENLANDICA Lehman, 1959 (Fig. 3) (Forey, 1998). The ventral margin of the anterior part of the lower jaw bears Referred Specimen-Mus6um national d'Histoire naturelle, a distinct crest. This ridge delineates the ventral border of a Paris, no. MNHN ARD 258, isolated left lower jaw. Strud, well-marked depression, identified here as the labial pit (Fig. 3). Namur Province, Belgium, upper middle Famennian, Upper De- This depression is composed entirely of dermal bone, and there vonian. is no trace of Meckelian bone on the specimen. The anterior part

This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions 282 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 26, NO. 2, 2006

Famennian of Belgium (Cldment et al., 2004). Soederberghia is Friedman,M. 2003a. New elements of the Devonian lungfishSoeder- the only cosmopolitan Famennian dipnoan from the Old Red berghiagroenlandica (Sarcopterygii, Dipnoi), with commentson the the MPhil Uni- Sandstone Continent, as is Holoptychius among porolepiformes, status of Rhynchodipteridae.Unpublished Thesis, of 89 Eusthenodon among tristichopterids, or Groenlandaspis, versity Cambridge, pp. Friedman,M. 2003b.New insightson the Late Devonian lungfishSoe- Remigolepis, and Bothriolepis among placoderms (Lebedev, derberghiagroenlandica (Sarcopterygii: Dipnoi) from East Green- 2004). land with commentson the status of Rhynchodipteridae;in H.-P. Schultze,E. Luksevics,and D. Unwin (eds.), The GrossSymposium ACKNOWLEDGMENTS 2: Advancesin Palaeoichthyology.Ichthyolith Issues SpecialPubli- cation 7:22-23. We are indebted to E. and to M.-C. Poty (Liege University) Z. 1950.Les dipneustes du massifde Ste Groessens Van Catholic Gorizdro-Kulczycka, Ddvoniens Dyck (Louvain-la-Neuve University) Croix.Acta GeologicaPolonica 1:53-105. for loan of historical specimens such as the 'oldest known am- Graham-Smith,W., and T. S. Westoll. 1937.On a new long-headeddip- phibian', the isolated entopterygoid, and the isolated vertebrae. noan fish from the Upper Devonian of ScaumenacBay, P. Q., We would like to thank J. Thorez and M. Streel (Liege Univer- Canada. Transactions of the Royal Society of Edinburgh 59: sity) for help and support, and for comments on the sedimentol- 241-268. ogy and dating of the Famennian outcrops of Belgium. We thank Gross, W. 1956.Uber Crossopterygierund Dipnoer aus dem baltischen Oberdevonin einer all others members of the Belgian-French-Canadian-Swedish Zusammenhang vergleichendenUntersuchung des Porenkanalsystems palaozoischer Agnathen und Fische. field mission, i.e., P. Janvier, V. Dupret, A. Blieck, C. Derycke- Khatir, L. Pille, M. and D. as well as our three KungligaSvenska VetenskapsAkademiens Handlingar 2(5):1-140. Brazeau, Snitting, Janvier,P., and Clement,G. 2005. A new groenlandaspididarthrodire associated research Histoire de la groups: Ddpartement Terre, (Vertebrata:Placodermi) from the Famennianof Belgium. Geo- national Musdum d'Histoire naturelle, Paris, France; Ddparte- logica Belgica 8(1-2):51-67. ment de Gdologie, Universitd de Liege, Belgium; and USTL Jarvik,E. 1952.On the fish-liketail in the ichthyostegidstegocephalians Sciences de la Terre, Lille, France. Thanks are extended to M. withdescriptions of a new stegocephalianand a new crossopterygian Friedman, P. Janvier, and M. Brazeau for their useful comments from the Upper Devonian of East Greenland. Meddelelser om on the manuscript. We express our sincere thanks to the staff of Gronland114(12):1-90. the Communal Administration of Gesves, Namur Province, and Jarvik,E. 1967. On the structureof the lower jaw in dipnoans:with a to P. Mahoux, M. D. S. A. V. P. descriptionof an early Devonian dipnoan from Canada,Melano- especially Evrard, Bruaux, Thiry, et nov. Journalof the B. and to J.-P. Lefebvre for their and use- gnathuscanadensis gen. sp. LinneanSociety Beaujeant, important 47:155-183. ful in the of the field missions in (Zoology) support logistic part Belgium. Jarvik,E. 1980. Basic Structureand Evolution of Vertebrates,Vol. 1. This is ongoing project supported by the Swedish Research AcademicPress, London, 575 pp. Council through a grant to P. E. Ahlberg. Johanson,Z., and A. Ritchie2000. A new late Famennianlungfish from *This paper is a contribution to IGCP Project 491 (Middle New South Wales, Australia,and its bearingon Australian-Asian Palaeozoic Vertebrate Biogeography, Palaeogeography, and Cli- terranerelations. Alcheringa 24:99-118. mate). Krupina,N. I. 1987.A new dipnoanfish fromthe UpperDevonian of the Tula Region. PaleontologicalJournal 3:37-43. LITERATURE CITED Krupina,N. I. 1999.A new dipnoanfrom the Upper Devonian locality Rybnitsain Oryol Region. PaleontologicalJournal 33:627-629. Ahlberg, P. E. 1991. A re-examinationof sarcopterygianinterrelation- Lebedev,O. A. 2004.A new tetrapodJakubsonia livnensis from the early ships,with special reference to the Porolepiformes.Zoological Jour- Famennian(Devonian) of Russia and palaeoecologicalremarks on nal of the LinneanSociety 103:241-287. the Late Devonian tetrapodhabitats. Acta UniversitatisLatviensis Ahlberg,P. E., and N. H. Trewin.1995. The postcranialskeleton of the 679:79-98. Middle Devonian lungfishDipterus valenciennesi.Transactions of Lehman,J.-P. 1955. Les Dipneustesdu Ddvonien sup6rieurdu Groen- the Royal Society of Edinburgh85:159-175. land. Comptes-rendusdes S6ances de l'Acaddmie des Sciences, Ahlberg, P. E., Z. Johanson,and E. B. Daeschler.2001. The Late De- Paris 240:995-997. vonianlungfish Soederberghia (Sarcopterygii, Dipnoi) fromAustra- Lehman,J.-P. 1959. Les Dipneustesdu D6vonien sup6rieurdu Groen- lia and NorthAmerica, and its biogeographicalimplications. Journal land. Meddelelserom Gr0nland160:1-58. of VertebratePaleontology 21:1-12. Leriche,M. 1931.Les PoissonsFamenniens de la Belgique.Mdmoires de Berg, L. S. 1940. Classificationof fishes, both recent and fossil. Trudy la Classe des Sciences de l'Acad6mie Royale de Belgique 10(5): ZoologicheskogoInstituta, Akademiya Nauk SSSR, Leningrad5: 1-72. 1-141. Lohest,M. 1888a.Recherches sur les poissonsdes terrainspaldozoiques Campbell,K. S. W., and R. E. Barwick.1987. Paleozoic lungfishes--a de Belgique. Poissons des Psammites du Condroz, Famennien review.Journal of MorphologySupplement 1:93-131. superieur.Annales de la SocietdGdologique de Belgique,M6moires Campbell,K. S. W., and R. E. Barwick.1988. Uranolophus: a reappraisal 15:112-203. of a primitivedipnoan. Memoirs of the Associationof Australasian Lohest, M. 1888b. D6couvertedu plus ancien amphibienconnu et de Palaeontologists7:87-144. quelques fossiles remarquablesdans le Famennien sup6rieurde Campbell,K. S. W., and R. E. Barwick.1990. Paleozoic dipnoan phy- Modave. Annales de la Soci~t6 Gdologique de Belgique:129-137. logeny:functional complexes and evolutionwithout parsimony. Pa- Long, J. A. 1992. Cranial anatomy of two new Late Devonian Lungfishes leobiology 16:143-169. (Pisces: Dipnoi) from Mount Howitt, Victoria. Records of the Aus- Campbell,K. S. W., and R. E. Barwick. 2002. The axial postcranial tralian Museum 44:299-318. structureof Griphognathuswhitei from the Upper Devonian Gogo Miles, R. S. 1977. Dipnoan (lungfish) skulls and the relationships of the Formationof WesternAustralia: comparisons with other Devonian group: a study based on new species from the Devonian of Australia. dipnoans.Records of the WesternAustralian Museum 21:167-201. Zoological Journal of the Linnean Society 61:1-328. Campbell,K. S. W., and M. W. Bell. 1982.Soederberghia (Dipnoi) from Moy-Thomas, J. A. 1939. Palaeozoic Fishes. Methuen, London, 149 pp. the Late Devonian of New South Wales. Alcheringa6:143-150. Miller, J. 1845. Uber den Bau und die Grenzen der Ganoiden, und iber Cldment,G., P. E. Ahlberg,A. Blieck, H. Blom, J. A. Clack,E. Poty, J. das nattirliche System der Fische. Physikalisch-mathematische Ab- Thorez,and P. Janvier.2004. Devonian tetrapodfrom WesternEu- handlungen der kiniglichen Akademie der Wissenschaften zu Ber- rope. Nature 427:412-413. lin 1845:117-216. Cloutier, R. 1996. Dipnoi (Akinetia: Sarcopterygii); pp. 198-226 in H.-P. Panchen, A. L. 1970. Batrachosauria (Anthracosauria); in O. Kuhn (ed.), Schultze and R. Cloutier (eds.), Devonian Fishes and Plants of Handbuch der Palioherpetologie, part 5A. Gustav Fischer, Stutt- Miguasha, Quebec, Canada. Verlag Dr. Friedrich Pfeil, Miinchen. gart, 83 pp. Forey, P. L. 1998. History of the Coelacanth Fishes. Chapman and Hall, Pridmore, P. A., and R. E. Barwick. 1993. Post-cranial morphologies of London, 419 pp. the Late Devonian dipnoans Griphognathus and Chirodipterus and

This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions CLEMENT and BOISVERT-DEVONIAN LUNGFISH FROM BELGIUM 283

locomotor implications. Memoirs of the Association of Australasian Schultze, H.-P. 1970. Die Histologie der Wirbelkirper der Dipnoer. Palaeontologists 15:161-182. Neues Jahrbuch ftir Geologie und Paliontologie, Monatshefte 135: Pridmore, P. A., K. S. W. Campbell, and Barwick, R. E. 1994. Morphol- 311-336. and ogy phylogenetic position of the holodipteran dipnoans of the Schultze, H.-P. 1992. A new long-headed dipnoan (Osteichthyes) from Upper Devonian Gogo Formation of nortwestern Australia. Philo- the Middle Devonian of Iowa, USA. Journal of Vertebrate Paleon- sophical Translations of the Royal Society of London B 344: tology 12:42-58. 105-164. Schultze, H.-P. 1993. Osteichthyes: Sarcopterygii; pp. 657-663 in M. J. Romer, A. S. 1955. or Herpetichthyes, Amphibioidei, Choanichthyes Benton (ed.), The Fossil Record 2. Chapman and Hall, London. Sarcopterygii? Nature 176:126. Schultze, H.-P. 2001. Melanognathus, a primitive dipnoan from the Rosen, D. E., P. L. Forey, B. G. Gardiner, and C. Patterson. 1981. Lung- Lower Devonian of the Canadian Arctic and the interrelationships fishes, tetrapods, and Bulletin of the paleontology, plesiomorphy. of Devonian Journal of Vertebrate Paleontology 21: American Museum of dipnoans. Natural History 167:159-276. 781-794. Sive-Siderbergh, G. 1937. On Rhynchodipterus elginensis n. g., n. sp., Young, G. C. 1999. Preliminary report on the biostratigraphy of new representing a new group of dipnoan-like Choanata from the Upper placoderm discoveries in the Hervey Group (Upper Devonian) of Devonian of East Greenland and Scotland. Arkiv for Zoologi 29: 1-8. central New South Wales. Records of the Western Australian Mu- seum 57:139-150. Schultze, H.-P. 1969. Griphognathus Gross, ein langschnauziger Dipnoer Supplement aus dem Oberdevon von Bergisch-Gladbach (Rheinisches Schiefer- gebirge) und von Lettland. Geologica et Palaeontologica 3:21-79. Submitted 1 August 2005; accepted 6 September 2005.

This content downloaded from 157.193.10.229 on Tue, 07 Jul 2015 14:17:10 UTC All use subject to JSTOR Terms and Conditions