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Species Glyptoxoceras rugatum Forbes 1846

Diagnosis / Definition:
Kennedy & Henderson (1992):
Description: The early growth stages are shown by a series of evenly ribbed and occasionally constricted fragments down to a whorl heights of as little as 3mm. They vary from compressed to subcircular in cross section and are slightly curved, defining obtuse or acute angels, or straight. The same shapes characterize the largest fragments, indicating a planispiral shell, polygonal in outline, and made up of straight shafts connected by obtuse or acute angels. The lectotype of Hamites rugatus (Forbes, 1846, pl. 11, fig. 2) is a wholly septate fragment consisting of two diverging shafts linked by a curved sector (Pl. 1, figs. 1-2). It has a maximum preserved whorl height of 198 mm, and a whorl breadth to height ratio of 0.87; the rib index is 8, the ribs blunt, somewhat weakened on the dorsum but strengthening progressively across the flanks, and strong and transverse on the venter. They are prorsiradiate at the smallest and largest diameters preserved, but markedly rursiradiate on the curved sector. The lectotype of Hamites subcompressus Forbes, 1846 (Pl. 1, figs. 13-16) is a curved body chamber, 150 mm long. The whorl section is compressed oval, with flattened subparallel flanks and a broadly rounded dorsum and venter. The costal whorl breadth to height ratio is 0.73, and the rib index varies between 7 and 8. The ribs are relatively weak and transverse on the dorsum, but strenghten across the flanks, where they are feebly prorsiradiate, and on the venter, where they are transverse. The ribs are distinctly narrower than the interspaces, but show some variation, being blunter on the internal mould than where shell is preserved. The holotype of Glyptoxoceras circulare Shimizu, 1935 (Text-figs. 10-11; Pl. 4, fig. 12) is a broken, straight shaft 95 mm long, with a circular cross section, embedded in matrix. The style of ribbing is, so far as visible, like that of the lectotype of subcompressus, with a rib index of six. Suture (Text-fig. 1A, E) with moderately incised, bifid lobes and saddles.
Ward & Kennedy (1993):
Description. - Specimens are crushed fragments, either straight shafts or shafts associated with curved sections. Greatest preserved whorl height 18 mm. Rib index 6-7. Ribs weak on dorsum, strengthening over dorsolateral area and flank to reach maximum development over venter. They are prorsiradiate, straight, and narrower than interspaces. Sutures not seen.
Discussion / Comments:
Kennedy & Henderson (1992):
Types: Lectotype, here designated, is BMNH C51110, the original of Forbes, 1846 (pl. 11, fig. 2), GSC R10499. The lectotype, here designated, of Hamites subcompressus Forbes, 1846, is shown in Plate 1, figures 12-16. it is broken into three parts (Pl. 1, fig. 16). The middle fragment (A) as noted by Philipps (1977, p. 125), GSC R10491, is the original of Kossmat (1895, pl. 19(5), fig. 10), and possibly the original of Forbes (1846, pl. 11, fig. 6). The adapical fragment (B) corresponds with Forbe's plate 11, figure 4a, c, the figured syntype of Hamites indicus. Fragment C was never figured. The lectotype, here designated, of Hamites nereis Forbes, 1846, plate 10, figure 7, is BMNH C51109, GSC R10502. In each case lectotypes are designated because Forbes cited the 'dimension of largest specimen', indicating that he possessed more than one of each species. The holotype of Glyptoxoceras circulare Shimizu, 1935 (text-figs. 10-11) is BMNH C51112. Possible paralectotypes of Hamites subcompressus are BMNH C51112, the original of Kossmat (1895, pl. 19(5), fig. 4c = GSC R10497; Pl. 2, fig. 16), and BMNH C 51103, the original of Kossmat (1985, pl. 19(5), fig. 11; Pl. 1, fig. 9, herein). Topotypes BMNH C51125 (Pl. 1, fig. 5), C51131 and C51139 are possible syntypes of Hamites indicus that belong to the present species, as are BMNH C51120-51122 (Pl. 1, figs. 10-11; Pl. 4, fig. 2), syntypes of Hamites undulatus Forbes, 1846; and BMNH C51096-51099, which are syntypes of Hamites largesulcatus Forbes, 1846 (Pl. 3, figs. 1-3; Pl. 4, figs. 13-15)(all ex Kaye and Cunliffe Collection). Other topotypes are BMNH C4049 (5 fragments, no history), C4050 (12 fragments, no history); C4109 (history uncertain, the original of Shimizu, 1935, figs. 6-9), C5114-5116 (no history), BMNH C24201 (4 fragments, ex Kaye Collection), C51126-51129 (ex Geological Society Collection), and C2402 (ex Kaye Collection). All are from the Valudavur Formation of Pondicherry, South India. Discussion: We regard Hamites rugatus, H. nereis and H. subcompressus of Forbes (1846) as synonyms, and as first revising authors select the name rugatus for the species. Also regarded as conspecific is G. circulare, and some of the syntypes of Hamites indicus of Forbes. The characteristic features of the species are the polygonal coiling, relatively large size, and rib density. When compared with other species in the Ponicherry fauna. Glyptoxoceras tenuisulcatum (Forbes, 1846) has an initial helix, and much finer ribbing that is oblique on the venter of the shaft, rather than being transverse (Pl. 2, figs. 2, 6, 8, 30). Glyptoxoceras largesulcatum (Forbes, 1846)(Pl. 3, figs. 4-9; Pl. 4, figs. 16-18) has very distant annular ribs, and is only known as straight shafts. G. indicum (Forbes, 1846) (Pl. 1, figs. 1, 3-5, 7, 9, 12-13) has a quite different coiling, with an initial helix, and oval planispiral later whorls, being adult at much smaller size (Pl. 1, figs. 3-4). When these species are removed from the Pondicherry fauna, the numerous remaining fragments from a species that is variable in whorl section and ribbing density, but of similar coiling and ribbing style. Hamites nereis (Forbes, 1846) (Pl. 2, figs. 10-11), referred to Pseudoxybeloceras by some authors, is a pathological rugatum in which ventral ribbing is interrupted, as a result of non-lethal injury to the mantle margin (Pl. 2, fig. 10). Of species from Pondicherry illustrated by d'Orbigny (1847a), one of the specimens of Hamites acuticostatus (pl. 3, figs. 11, 12) may belong here; it is shown as being 40 mm long, with a compressed whorl section, and rib index of 4. The second specimen illustrated by d'Orbigny (pl. 3, figs. 9-10) seems to be a G. largesulcatum. The Hamites simplex of d'Orbigny from Pondicherry (pl. 3, figs. 15-16) is a curved fragment 30 mm long, and is clearly G. rugatum. We refer a number of non-Indian records to G. rugatum on the basis of coiling style, ribbing and age (see synonymy), but reject small fragments that are significantly older where the coiling style is unclear, believing them to be unidentifiable as to genus in some cases. Thus fragments from the Upper Maastrichtian of the Maastricht area, described as Hamites rotundus by Binkhorst (1861) are referred to G. rugatum (see also Kennedy, 1987), as is G. recticostatus (Seunes, 1890) from the Upper Maastrichtian of southeastern France, and G. braziliense (Maury, 1930), from Brazil. Study of more than 100 Glyptoxoceras from the Upper Maastrichtian Miria Formation of western Australia (to be described elsewhere), has convinced us that G. indicum, G. circulare, G. nipponicum, and G. bullarense, Neohamites giralensis, N. rugatus, N. cardabiensis, N. largesulcatus, and N. soufoulisi of Brunnschweiler (1966) represent but a single form, conspecific with Glyptoxoceras rugatum. The Glyptoxoceras subcompressum of Ward (1976, p. 456, pl. 1, figs. 1-5; text-fig. 3) and Ward and Westermann (1976, text-figs. 1-3) has a quite different, ellipsoidal coiling when compared to the Pondicherry material, with an initial helix with its axis of coiling at 90° to that of the planispiral whorls; it is of late Santonian or early Campanian age. Occurrence: Maastrichtian of south India, Brazil, Chile (?), Western Australia, northern Spain, south-east France, and the Maastricht area in the Netherlands and Belgium.
Ward & Kennedy (1993):
Type.- This species is based upon a series of fragments from the Valudavur Formation of Pondicheny, south India, in the collections of the Natural History Museum, London, and lectotype designation is deferred until this material is published (Kennedy and Henderson, in prep.). Discussion.- These crushed fragments provide only rib density, style, and direction as clues to specific identity, but a comparison with the largest Indian example of Glyptoxoceras rugatum in the Forbes Collection reveals no obvious differences. Glyptoxoceras largesulcatum (Forbes, 1 846) (p. 1 1 7, Pl. 11, fig. 1; see Fig. 18.13) is easily differentiated by the low rib index (4.5) and large size. Glyptoxoceras tenuisulcatum (Forbes, 1846) (p. 116, Pl. 11, fig. 3) is in contrast densely and finely ribbed, with a circular cross section. Glvptoxoceras indicum (Forbes, 1846) p. 1 16, Pl. 1 1, fig. 4) (see Matsumoto, 1959, p. 167, Pl. 4 1, figs. 2-6, text-fig. 80) has a different coiling mode, seemingly lacking the long straight to slightly curved shafts of G. rugatum. with a circular whorl section. 0ccurrence.- In the Biscay sections this taxon ranges from the upper part of Member I (lower Maastrichtian) to the upper half of Member IV (upper Maastrichtian). The taxon shows a discontinuous range across the basin; at Zumaya it is one of the commonest ammonites, whereas it is rare or absent at the other Biscay sections. Immel, Klinger, and Wiedmann (1982) referred fragments from as low as the lower Santonian to the present species, but it is perhaps questionable how many of these generalized diplomoceratid scraps are conspecific. The most convincing records are Maastrichtian. The recorded geographic range from this stage is southeastern France, Spain, the Netherlands, south India, and western Australia.

 Ordo Ammonoidea
  Subordo Ancyloceratina
   Superfamilia Turrilitaceae
    Familia Diplomoceratidae
     Subfamilia Diplomoceratinae
      Genus Glyptoxoceras
       Species Glyptoxoceras rugatum

  Subordo Ancyloceratina
   Superfamilia Turrilitaceae
    Familia Diplomoceratidae
     Subfamilia Diplomoceratinae
      Genus Diplomoceras
       Species Glyptoxoceras rugatum
Synonym list:
Kennedy & Henderson (1992):
1846 Hamites subcompressus Forbes. - Forbes : p.116 pl. 11; fig. 6
1846 Hamites rugatus Forbes. - Forbes : p.117 pl. 11; fig. 2
1846 Hamites nereis Forbes. - Forbes : p.117 pl. 10, fig. 7
1847 Hamites acuticostatus d'Orbigny. - d'Orbigny : pl. 3; fig. 11-12 only
1847 Hamites simplex d'Orbigny. - d'Orbigny : pl. 3; fig. 15-17 (non d'Orbigny, 1842)
1850 Hamites subcompressus Forbes. - d'Orbigny : p.216
1850 Hamites rugatus Forbes. - d'Orbigny : p.216
1850 Hamites nereis Forbes. - d'Orbigny : p.216
1861 Hamites rotundus Sowerby?. - Binkhorst : p.34 pl. 5b, fig. 2-4; pl. 5c, fig. 1
1866 Anisoceras rugatum Forbes. - Stoliczka : p.178 pl. 85; fig. 10-13
1866 Anisoceras subcompressum Forbes. - Stoliczka : p.179 pl. 85; fig. 7
p 1866 Anisoceras indicum Forbes. - Stoliczka : p.181 pl. 85; fig. 3-5 only
? 1866 Anisoceras nereis Forbes. - Stoliczka : p.182 pl. 85; fig. 17-18
? 1866 Anisoceras sp. ?. - Stoliczka : p.179 pl. 85; fig. 19
non 1871 Anisoceras rugatum Forbes. - GRIESBACH : p.63 pl. 3; fig. 4
1890 Hamites recticostratus Seunes. - Seunes : p.239 pl. 9; fig. 6
1895 Hamites (Anisoceras) subcompressus Forbes. - Kossmat : 145 (49) pl. 19 (5); fig. 10-12
p 1895 Hamites (Anisoceras) rugatus Forbes. - Kossmat : p.146 [non pl. 19(5), fig. 7-9; = G. indicum, D. cylindraceum]
? 1895 Hamites (Anisoceras) sp. . - Kossmat : 148(52)
1895 Hamites (Anisoceras) nereis Forbes. - Kossmat : 148(52)
non 1903 Anisoceras subcompressum Forbes. - Whiteaves : p.338 pl. 45; fig. 1
non 1906 Hamites (Anisoceras) subcompressus Forbes. - WOODS : p.339 pl. 43; fig. 2
? 1925 Glyptoxoceras cf. rugatum Forbes. - Spath : p.30 pl. 1; fig. 4
? 1930 Diplomoceras (Hamites) sp. . - Wetzel : p.90
1930 Glyptoxoceras braziliense Maury. - Maury : p.184 pl. 11; fig. 6
1935 Glyptoxoceras rugatum Forbes. - SHIMIZU : p.273 text-fig. 1-9
1935 Glyptoxoceras circulare Shimizu. - SHIMIZU : p.272 text-fig. 10-11
1935 Glyptoxoceras subcompressum Forbes. - SHIMIZU : p.272 text-fig. 12
1935 Glyptoxoceras indicum var. intermedium Shimizu. - SHIMIZU : p.272
1940 Glyptoxoceras cf. rugatum Forbes. - Spath : p.47 pl. 1; fig. 1
non 1952 Diplomoceras? subcompressum Forbes. - USHER : p.110 pl. 29; fig. 3
1959 Glyptoxoceras rugatum Forbes. - Matsumoto : p.169
1959 Glyptoxoceras subcompressum Forbes. - Matsumoto : p.169
1962 Diplomoceras (Glyptoxoceras) cf. subcompressum Forbes. - Wiedmann : p.208 pl. 12; fig. 1-2
1966 Glyptoxoceras indicum Forbes. - BRUNNSCHWEILER : p.44 pl. 6, fig. 1-3; text-fig. 28
1966 Glyptoxoceras circulare Shimizu. - BRUNNSCHWEILER : p.46 pl. 6, fig. 4-6; text-fig. 29
1966 Glyptoxoceras nipponicum Shimizu. - BRUNNSCHWEILER : p.46 pl. 6, fig. 7-9; text-fig. 30
1966 Glyptoxoceras bullarense Brunnschweiler. - BRUNNSCHWEILER : p.47 pl. 6, fig. 10; text-fig. 31
1966 Neohamites giralensis Brunnschweiler. - BRUNNSCHWEILER : p.48 pl. 7, fig. 1-2; text-fig. 32
1966 Neohamites rugatus Forbes. - BRUNNSCHWEILER : p.49 pl. 7, fig. 4-6; text-fig. 33
1966 Neohamites cardabiensis Brunnschweiler. - BRUNNSCHWEILER : p.51 pl. 7, fig. 7-9; text-fig. 35
1966 Neohamites largesulcatus Forbes. - BRUNNSCHWEILER : p.51 pl. 1, fig. 8; pl. 8, fig. 3-6; text-fig. 36
1966 Neohamites soufoulisi Brunnschweiler. - BRUNNSCHWEILER : p.53 pl. 8, fig. 2; text-fig. 37
non 1966 Diplomoceras aff. subcompressum Forbes. - COLLIGNON : p.6 pl. 457; fig. 1861
non 1969 Glyptoxoceras ssp. subcompressum coarctum Collignon. - COLLIGNON : p.41 pl. 529; fig. 2084-2085
non 1976 Glyptoxoceras subcompressum Forbes. - Ward : p.456 pl. 1, fig. 1-5; text-fig. 3
non 1976 Glyptoxoceras subcompressum Forbes. - Ward & WESTERMANN : p.357 text-fig. 1-3
non 1976 Diplomoceras (Glyptoxoceras) subcompressum Forbes. - KLINGER : p.80 pl. 34; fig. 6
1977 Neohamites subcompressus Kennedy. - Kennedy : text-fig. 31.6
non 1982 Diplomoceras (Glyptoxoceras) subcompressum Forbes. - Immel et al. : p.26 pl. 9, fig. 4-5; pl. 10, fig. 7; pl. 11, fig. 4
1986 Glyptoxoceras cf. subcompressum Forbes. - Kennedy : text-fig. 9f-g
1986 Glyptoxoceras cf. circulare Shimizu. - Kennedy : text-fig. 9i-j
1987 Glyptoxoceras cf. circulare Shimizu. - Kennedy : p.180 pl. 4, fig. 1-3; pl. 26, fig. 7, 10-12, 15
1987 Glyptoxoceras cf. subcompressum Forbes. - Kennedy : p.179 pl. 26, fig. 1-6, 8-9, 13-14, 19-21
1992 Glyptoxoceras rugatum Forbes. - Kennedy & Henderson : p. 697, 701, 705, 71 pl. 1, fig. 1-2, 5-16; pl. 2, fig. 10-11, 14-29; pl. 3, fig. 1-3; pl. 4, fig. 2, 12-15; text-fig. 1A, E
Ward & Kennedy (1993):
1846 Hamites rugatus Forbes. - Forbes : p.117 pl. 11; fig. 2a, 2c
1992 Glyptoxoceras rugatum Forbes. - Henderson et al. : p.143 fig. 8-13 (with full synonymy)
1993 Glyptoxoceras rugatum Forbes. - Ward & Kennedy : p. 19, 50, 52 fig. 18.14, 43.10-43.12, 45.4
Stratigraphy - relative ages:
upper Maastrichtian - lower Maastrichtian: Ward & Kennedy (1993)
Maastrichtian: Kennedy & Henderson (1992)

Forbes,E.. (1846):
Report on the Cretaceous fossil invertebrates from southern India, collected by Mr. Kaye and Mr. Cunliffe . Transactions of the Geological Society of London Vol. 2(7) p. 97–174

d'Orbigny,A. (1847):
Paléontologie, Pls. 1-6 (Geologie Pls. 4-9).
In: M. de Dumont D'Urville, 1846-1 854, Voyage au Pole Sud et dans I'Oceanie sur les corvelles L'Astrolabe et la Zelée pendant les années 1837- 1838-1839-1840 sous le commandément de M. Dumont D'Urville Capitaine du Vaisseau. Pls. 1-9 Eds: Baudry, G..

d'Orbigny,A. (1850):
Prodrome de Paléontologie. Stratigraphique universelle des animaux mollusques & rayonnés faisant suitre au cours élémentaire de paléontologie et de géologie stratigraphique. , Cours Élémentaire de Paléontologie et de Géologie Stratigraphiques Vol. 2

Binkhorst,J.T.. (1861):
Monographie des gastropodes et des céphalopodes de la Craie Supérieure du Limburg.
In: Maastricht and Muller Frères Eds: Muquardt, G.. p. 17,83,44

Stoliczka,F.. (1866):
The fossil Cephalopoda of the Cretaceous rocks of southern India. Ammonitidae with revision of the Nautilidae etc. . Memoirs of the Geological Survey of India, (l), Palaeontologica Indica10-13 p. 155-216

GRIESBACH,C.L.. (1871):
On the geology of Natal in South Africa . Quarterly Journal ofthe Geological Society of London Vol. 27 p. 53-72

Seunes,J.. (1890):
Recherches géologiques sur les terrains secondaires et l' Eocene inferieur de la region sous-pyrénéene du sud-ouest de la France (Basses-Pyrenees et Landes). p. 250 pp.

Kossmat,F.. (1895):
Untersuchungen über die Südindische Kreideformation. Erster Theil . Beiträge zur Paläontologie Österreich-Ungarns und des Orient Vol. 9 p. 97-203

Whiteaves,J.F.. (1903):
On some additional fossils from the Vancouver Cretaceous, with a revised list of the species therefrom. . Geol. Surv. Can., Mesozoic Fossils Vol. 1(5) p. 309-409

WOODS,H.. (1906):
The Cretaceous fauna of Pondoland . Annals of the South African Museum Vol. 4 p. 275-350

Spath,L.F.. (1925):
On Senonian Ammonoidea from Jamaica . Geol. Mag. Vol. 62 p. 28-32

Maury,C.J.. (1930):
O Cretaceo da Parahyba do Norte. , Album das estampas p. 305

Wetzel,W.. (1930):
Die Quiriquina-Schichten als Sediment und paläontologisches Archiv . Palaeontographica Vol. 73 p. 49-104

SHIMIZU,S.. (1935):
The Upper Cretaceous cephalopodes of Japan, part 1. . Journal of the Shanghai Science Institute Vol. 2(2) p. 159-226

Spath,L.F.. (1940):
On Upper Cretaceous (Maastrichtian) Ammonoidea from Western Australia . Jl. R. Soc. West. Aust. Vol. 26 p. 41-57

USHER,J.L.. (1952):
Ammonite faunas of the Upper Cretaceous rocks of Vancouver Island . Bulletin of the Geological Survey of Canada Vol. 21 p. 1-182

Matsumoto,T.. (1959):
Upper Cretaceous ammonites of California, Part II. . Memoirs of the Faculty of Science, Kyushu University, Series D, Geology, Special Volume Vol. 1 p. 1-172

Wiedmann,J.. (1962):
Ammoniten aus der Vascogotischen Kreide (Nordspanien). 1, Phylloceratina, Lytoceratina. . PalaeontographicaA118 p. 119-237

Upper Cretaceous ammonites from the Carnarvon Basin of Western Australia 1: the heteromorph Lytoceratina. . Bulletin of the Bureau of Mineral Resources, Geology and Geophysics Vol. 58 p. 5-58

COLLIGNON,M.. (1966):
Atlas de fossiles caracteristiques de Madagasca (Ammonites). , XIV, Santonien p. X+134 (455-513)

COLLIGNON,M.. (1969):
Atlas des fossiles caracteristiques de Madagascar (Ammonites). (Campanien inferieur). Vol. 15 p. 216 pp

Ward,P.D.. (1976):
Upper Cretacheous ammonites (Santonian-Campanian) from Orcas Island, Washington . J. Palaeont. Vol. 50 p. 454-461

Ward,P.D.. and WESTERMANN,G.E.G.. (1976):
Sutural inversion in a heteromorph ammonite and its implications for septal formation . Lethaia Vol. 9 p. 357-361

KLINGER,H.C.. (1976):
Cretaceous heteromorph ammonites from Zululand . Memoirs of the Geological Survey of the Republic of South Africa Vol. 69 p. 142 pp

Kennedy,W.J.. (1977):
Ammonite evolution.
In: Patterns of Evolution Eds: Hallam, A.. p. 251-330

Immel,H..; KLINGER,H.C.. and Wiedmann,J.. (1982):
Die Cephalopoden des Unteren Santon der Gosau von Brandenburg/Tirol, Österreich . Zitteliana Vol. 8 p. 3-32 (11pls)

Kennedy,J.W.. (1986):
The Campanian-Maastrichtian ammonite sequence in the environs of Maastricht (Limburg, the Netherlands) . Newsletters on Stratigraphy Vol. 16 p. 149-168

Kennedy,W.J.. (1987):
The ammonite faunas of the type Maastrichtian, with a revision of Ammonites colligatus Binkhorst, 1861 . Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre Vol. 56 p. 151-267

Henderson,R.A..; Kennedy,W.J.. and McNamara,K.J.. (1992):
Maastrichtian heteromorph ammonites from the Carnarvon Basin, Western Australia . Alcheringa Vol. 16 p. 133-170

Kennedy,W.J.. and Henderson,R.A.. (1992):
Heteromorph ammonites from the Upper Maastrichtian of Pondicherry, south India . Palaeontology Vol. 35 p. 693-731

Ward,P.D.. and Kennedy,W.J.. (1993):
Maastrichtian Ammonites from the Biscay Region (France, Spain) . Memoir (The Paleontological Society), Journal of Paleontology34 (Supplement to Vol. 67)(5) p. 1-58

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