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This Content Downloaded from 157.193.10.229 on Tue, 07 Jul 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).
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