Parengonoceras bassei ( BATALLER , 1954)

Latil, Ouvillé Ataller Jean-Louis & Aly, Mohamed Fouad, 2012, Knemiceras gracile D, 1916: a misunderstood Early Albian ammonite from north Sinai (Egypt), and considerations on the genus Platiknemiceras B, 1954, Revue de Paléobiologie 31 (1), pp. 159-170 : 162-166

Parengonoceras bassei ( BATALLER, 1954)

Fig. 4

1949. Knemiceras compressum HYATT. - DUBOURDIEU, p. 26, figs 1, 2.

1954. Knemiceras (Platiknemiceras) bassei BATALLER , p.175, fig. 1.

1956. Knemiceras compressum HYATT. - DUBOURDIEU, p. 150.

1959. Platyknemiceras bassei BATALLER , p. 65, fig. 826.

1961. Platiknemiceras bassei BATALLER. - CASEY, p. 355, fig. 1a-b.

1979. Knemiceras hachouri MASSE & THIEULOY , p. 68.

? 1982. Parengonoceras hachourii ( DUBOURDIEU, 1953) .- RENZ, p. 31, pl. 3, fig. 4; text fig. 19c-d.

1992. Platiknemiceras bassei BATALLER. - CAZALDA & URQUIOLA, p. 74, 1 fig.

1994. Platiknemiceras bassei BATALLER. - MARTINEZ, GRAUGES & SALAS, pl. 3, fig. 4.

1996. Platiknemiceras bassei BATALLER, 1954 .- WRIGHT, CALLOMAN & HOWARTH, p. 130, fig.100: 5a (not 5b).

1996. Platiknemiceras sp. - WRIGHT, CALLOMAN & HOWARTH, p. 130, fig.100: 5c.

? 2010. Platiknemiceras aff. bassei BATALLER, 1954 .- BULOT, p. 171, pl. 1, fig. 1.

2010. Platiknemiceras bassei BATALLER, 1954 .- BUJTOR, p. 13, fig.10, only the lateral view.

2011. Parengonoceras bassei ( BATALLER, 1954) .- LATIL, p. 349, fig. 24; pl. 3, figs 37-41; pl. 4, figs 1-6.

Type material: The lectotype ( MGSB.11478), designated by CASEY (1961, p. 23), has been figured by BATALLER (1954, p. 175, figs 1, 2); BATALLER (1959, p. 65, fig. 826); CAZALDA & URQUIOLA (1992, p. 74, photo on page 75), herein figured ( Fig. 4).

Dimensions of the lectotype:

D: 60 (100) – Wh: 32 (53) – Wb: 17 (28) – U: 6 (10) – v: 3 (5) – Wb/Wh: 0.53

Description: The species was described on the basis of two specimens identified from the Early Albian of NE Spain (Tarragona). The lectotype is preserved as a calcareous internal mould of a non-adult phragmocone. The coiling is involute, with a narrow umbilicus comprising 10% of the diameter. The umbilical wall is shallow and almost vertical. The whorl section is subtriangular, compressed, with a maximum width at the umbilical seam. The flanks are feebly convex, converging to a narrow, bicarinate, slightly concave venter. Strongly, prorsiradiate, faint ribs arise to umbilical seam, tending to blunt on the inner third of the flanks. The ribs curve backward at mid flanks, becoming radial on the outer third of the flanks and tending to be projected outward near the ventrolateral edge. The ribs, becoming fainter at the end of the preserved last whorl, are replaced by growth striae. Both the body chamber and the adult growth stages are unknown. The knemiceratid suture line is diagnostic, with broad first lateral lobe, frilled lobes and indentated saddles. This species seems to show a low rate of growth, with short distance between two sutures, even on the internal whorls. MARTINEZ et al. (1994, pl. 3, fig. 4) figured a specimen from Les Ventoses, Marmellà, Barcelona (MGSB 1044, probable paralectotype). This juvenile specimen, 28 mm in diameter, shows rather strong, prorsiradiate, sinuous riblets, rising on umbilical wall.

An additional fragment, MGSB 8689, a 180° sector of a phragmocone, about 80 mm in diameter, is characteristic of the species, with the ornament that tends to be effaced with age. The venter remains very narrow with age (v/D = 0,045), becoming slightly convex.

Discussion: This species represents the oldest known representatives of the genus Parengonoceras SPATH, 1924 . It resembles superficially Knemiceras gracile H. DOUVILLÉ, 1916 (p. 128, pl. 16, fig. 9), which has ventrolateral bullae, coarse and wide ribs on the flanks of the juvenile, and a suture with entire saddles.

Knemiceras deserti MAHMOUD, 1956 (p. 63, pl. 4, figs 3-5; text-figs 39-41) also shows a suture with entire saddles. These two last species show strong affinities with the genus Knemiceras s.str. Parengonoceras bassei is very similar in shape to Knemiceras hachourii DUBOURDIEU, 1953 (p. 23, pl. 2, fig. 10-18; pl. 3, fig. 1-5) and differs mainly by its grooved ventral area, the lack of true ventrolateral tubercles at a similar diameter.

Parengonoceras cf. hahourii (DUBOURDIEU) ( RENZ, 1982, p. 31, pl. 3, fig. 4; text-fig. 19 c-d) from the upper part of Machiques Formation (? transition Aptian- Albian), Quebrada Maraca, Venezuela, is characterized by a lack of lateral tuberculation, falcoid growth striae and bicarinate ventral area, and probably belongs to Parengonoceras bassei , differing only by its strongly frilled suture.

The specimen recently figured by BULOT (2010, pl. 1, fig. 1) as Platiknemiceras aff. bassei BATALLER, 1954 , from the Aptian-Albian transition of Kuh-e-Bangestan ( Iran) is too poorly preserved to allow a determination.

BUJTOR (2010) re-figured two drawings of CASEY (1961, fig. 1a, c) as “reproduction of the figure of BASSE (1954, p. 175)” (sic!). The lateral view corresponds to a drawing by CASEY (1961, fig. 1a) of the original figure of BATALLER (1954, fig. 1), while the ventral view is a drawing of a peruvian specimen, identified by CASEY (1961, fig. 1c) as Platiknemiceras sp. The same mistake is recorded in WRIGHT et al. (1996, p. 130).

Age: The type species Knemiceras (Platiknemiceras) bassei BATALLER, 1954 is known from Marmella, Montmell Formation (Salou-Garraf, Tarragona). No other ammonites co-occur with this species in its type locality. Findings in the laterally equivalent Escucha Formation, comprising “ Hypacanthoplites milletianus (d’ORBIGNY), Hypacanthoplites milletioides CASEY and Sonneratia subglabra CASEY ” (MARTINEZ, GRAUGES & SALAS, 1994; MORENO- BEDMAR et al., 2008), allow to assign an Early Albian age to Platiknemiceras bassei ( Leymeriella tardefurcata Zone to Sonneratia chalensis Zone ). MORENO- BEDMAR et al. (2008) mention co-occurrence of ‘ Hypacanthoplites ’ and Parengonoceras bassei within the same bed in the basal Escucha Formation (Puntarro de Traiguera section, Cuenca del Maestrat, E Cordillera Ibèrica). This strongly suggests a basal Early Albian age for Parengonoceras bassei .

Occurrence: Early Albian of Spain ( BATALLER, 1954), Tunisia ( LATIL, 2011) and maybe Venezuela ( RENZ, 1982).

IV. DISCUSSION ABOUT THE TAXONOMIC POSITION OF THE TAXA FORMELY REFERRED TO THE GENUS PLATIKNEMICERAS BATALLER, 1954

Iran: CASEY (1961, p. 354) reported three specimens from the Albian of Hamiran, south Iran ( SPATH, 1931, p. 339, text-fig 111f-h; CASEY, 1961, text-fig. 1c: suture lines). SPATH attributed these three specimens to Knemiceras of the group of attenuatum -gabbi (sensu lato). He interpretated them as probable new species, and considered them as passage-forms between Knemiceras and Engonoceras , following PERVINQUIÈRE’ s opinion (1907). These specimens are housed in the collections of the Natural History Museum of London under the numbers C68402, C68403 and C648406 ( Figs 5, 6). These specimens, represented by rather large-sized phragmocones, show a compressed whorl section, narrow umbilicus, slightly convergent flanks and a narrow, bicarinate venter. The ventrolateral edges are crenate to tuberculate, and ventrolateral tubercles, when present, are opposite on the venter. One specimen ( NHML.C68406, Fig. 6 d-f) has radially elongated, periumbilical swellings, from which arise single, slightly flexuous, narrow, coarse ribs that tend to be effaced with age. Morphological features of this material strongly suggest close affinities to the genus Knemiceras . Pending a revision of the south iranian engonoceratid faunas, and following SPATH’ s opinion, we herein consider this material as belonging to one or several undescribed new species.

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Lebanon: Knemiceras subcomplicatum BASSE, 1940, p. 434 , pl. 3, fig. 2, 3, from the Albian of Lebanon , is based on two specimens. The holotype ( MHNP. R052050 : pl. 3, fig. 2), from Bhamdoum, is a crushed, distorted and weathered fragment of an adult phragmocone, about 130 mm diameter, with a compressed whorl section, narrow umbilicus and narrow venter. No ornament can be observed because of weathering. According to BASSE, the main features of this species are based mainly on the suture line, which is supposed to be characterized by deep lobes, strongly frilled saddles and numerous accessory elements. The suture being strongly weathered, cannot be clearly studied. In addition, we agree with YOUNG’ s statement (1957, p. 2): “In my opinion, authors who designate genera and/or species on sutures only are daydreaming” .

The paratype ( MNHP. R 52057, pl. 3, fig. 3), from Maaser-ech-Chouf, is a complete specimen, preserved as a weathered, slightly distorted, calcareous internal mould, 115 mm diameter, comprising a 180° sector of body chamber. The whorl section is compressed, with a maximum width at about mid flanks. The umbilicus is narrow, with a shallow steep umbilical wall. The flanks are slightly convex, the venter is narrow and bicarinate, tending to become convex and wider on the end of body chamber. The ornament on ventrolateral edges is obscured, probably because of weathering. The inner part of the last whorl (end of the phragmocone) shows feeble, coarse ribs arising by two or three from coarse umbilical bullae. BASSE (1940, p. 434) mentions numerous tiny, ventrolateral tubercles on the phragmocone .

Knemiceras subcomplicatum BASSE, 1940 , is referred to the genus Platiknemiceras by CASEY (1961, p. 354) without justification. From the main features of the paratype, this species should be placed within the genus Knemiceras , but we are not even sure that the two specimens described by BASSE belong to the same species. The state of preservation of the holotype leads us to regard the present species as a nomen dubium.

Egypt: Knemiceras deserti MAHMOUD, 1956 (p. 63, pl. 4, fig. 3-5, text fig. 39-41), was described from the Early Albian of Gabal Manzour, north Sinai, Egypt (see also CASEY, 1961, p. 354), for small sized, compressed ammonites, with subtriangular whorl section, narrow umbilicus, steep umbilical wall, nearly smooth flanks, narrow venter, slightly crenate ventrolateral edges and faint ventrolateral bullae that are opposite on venter. This species is in need of a revision, but its suture suggests strong affinities with the genus Knemiceras . A careful examination of the paleontological collections in Geology Department of Cairo University (Egypt), has convinced us that Knemiceras deserti represents an extreme, compressed morphology of Knemiceras douvillei BASSE, 1940 .

Algeria: Engonoceras jullieni BASSE, 1940, p. 439 , textfig. 11; pl. 4, fig. 4, from the Albian of Khenchela, Algeria, is based on a poorly preserved, eroded and distorted specimen, from an unknown locality and stratigraphic level. This taxon is herein regarded as a nomen dubium.

GEYER (1995, p. 8), placed Placenticeras saadensis PERON, 1890, p. 19 ; pl. 16, figs 3-7, within the genus Platiknemiceras . BREISTROFFER (1940a and b) regarded this species as a Knemiceras . On the other hand, LATIL (1989) identified it as an Engonoceras . This Late Albian species from Algeria and Tunisia would be better placed within the genus Hypengonoceras SPATH, 1922 , as a result of its age and its sutural similarities.

Tunisia: CASEY (1961, p. 354), placed Knemiceras hachourii DUBOURDIEU, 1953 (p. 23, text-fig. 7; pl. 2, figs 10-18; pl. 3, figs 1-5) within the genus Platiknemiceras . LATIL (2011, p. 351) attributed this species to Parengonoceras .

Spain: Platiknemiceras valencianum MAS &WIEDMANN (1980, p. 265, figs 6, 7A, 8A) from La Ermita de Chera section, Valencia Province, Spain (Late Albian), is based on a poorly preserved fragment, but its suture shows strong affinities with sutures of Knemiceras sensu lato, such as Knemiceras uhligi ( CHOFFAT, 1886) . Furthermore, the ventrolateral clavi of Platiknemiceras valencianum are alternating on the ventral area. The present authors are satisfied to exclude this species from either Knemiceras or Platiknemiceras auctorum .

France: Platiknemiceras sequanense DESTOMBES, 1979, p. 114 , pl. 4, fig. 3, was described from a single specimen from Quarry West, Bois du Perchois, Aube, France, level 1b (Early Albian, Sonneratia dutempleana zone, Cleoniceras floridum Subzone of DESTOMBES). The holotype is a crushed and distorted fragment, represented by the end of the phragmocone and a 270° sector of body chamber. The body chamber has been crushed before fossilisation. The umbilicus is very narrow, with rather deep, rounded umbilical wall. The whorl section is subtriangular, with maximum width just above the umbilical seam. Then the flanks are convergent to a narrow, slightly concave bicarinate venter (v/D = 0.05). The author mentions smooth flanks, but an in-deep examination of the type reveals the presence of faint, flexuous growth striae on the body chamber. The suture is poorly preserved, showing numerous, frilled elements. From the same level, TOUCH & MATRION (1995, pl. 3, fig. 5) have figured an unregistered specimen identified as Platiknemiceras sp. This incomplete specimen, 55 mm in diameter, is characterized by a flat, narrow, bicarinate venter and strongly flexuous growth striae. MATRION (2010, p. 162, figs 116A-C) has figured two additional specimens from le Perchois quarries of Platiknemiceras sequanense . This species could be phylogenetically related to Parengonoceras bassei ( BATALLER, 1954) .

USA: Platiknemiceras flexuosum KENNEDY, LANDMAN & COBBAN, 1998, p. 38 , fig. 51, 52 ( Knemiceras sp. nov., CASEY, 1961, p. 355), from the Glen Rose Limestone (Early Albian) of Texas, is characterized by its strongly flexuous ribs, its convergent outer flanks and its narrow, slightly concave, bicarinate ventrer. This species is herein regarded as a primitive morphology of Texan engonoceratoids and is provisionally placed within the genus Parengonoceras . However, we believe that the north american engonoceratids most probably derive from some obscure Early Albian, tethyan stock, and have evolved in isolation, on the Texan platforms. In our opinion, this group deserves a separate generic status. It should be noted that this species and Parengonoceras roemeri ( CRAGIN, 1893) are contemporaneous.

Venezuela: Some discoidal specimens with a compressed whorl section and feeble ornament have been reported from the Early Albian of Venezuela ( RENZ, 1982).

Parengonoceras cf. hachourii (DUBOURDIEU) ( RENZ, 1982, p. 31, pl. 3, fig. 4; text-fig. 19c, d: specimen MB.Re2439), from the upper part of Machiques Formation in the Quebrada Maraca (Perijà river, Venezuela), characterized by the lack of lateral tuberculation, bicarinate ventral area, and prorsiradiate flexuous striae. It differs from Parengonoceras bassei mainly by its suture with strongly frilled saddles and somewhat less convex flanks. This ammonite was collected in the same stratigraphic levels of Prolyelliceras gevreyi ( JACOB, 1907) , providing an Early Albian age, Leymeriella tardefurcata Zone to base of Douvilleiceras mammillatum Superzone.

? Parengonoceras sp. ( RENZ, 1982, p. 32, pl. 4, fig. 2), from the top of Machiques Formation, Quebrada Macoita, Venezuela, is probably contemporaneous to Prolyelliceras gevreyi ( JACOB, 1907) . This specimen, with compressed section, umbilical tubercles and flexuous striae, is close to Parengonoceras caneroti COLLIGNON, 1981b , from which it differs by its strong ventrolateral clavi that are opposite on the ventral area.

Colombia: Platiknemiceras columbiana ETAYO- SERNA, 1979 from Cerro Bejucal, Colombia, basal part of the Capotes Member (?Middle Albian), is poorly figured (p. 76, pl. 11, fig. 7: holotype by monotypy C-169/ UNCMHNCPHt.N58). It is characterized by “A) finely crenate ventrolateral carinae, B) the trend of the bundles of lirae, radial on the central and adumbilical third of the flank, adorally concave on the adventral third” (ETAYO- SERNA, 1979, p. 76).

BREISTROFFER (1952, p. 2634) mentioned from the basal Middle Albian of Cundinamarca, Colombia, discoidal, compressed and involute specimens attributed to the genus Knemiceras , with a costation reduced to very fine striae and a narrow bicarinate venter.This material, placed by CASEY (1961, p. 355) in Platiknemiceras , has been borrowed by us from the collections of the Université Joseph Fourier (Grenoble, France). This material consists of two fragments from Quipilé, and an additional one from Viota, Quebrada Tasajera (Cundinamarca, Colombia). A small fragment of an internal mould of a juvenile specimen from Quipilé ( UJF. ID 10886), about 30 mm long, shows a narrow, flat, bicarinate venter. It has a compressed, subtriangular whorl section, with subparallel flanks ( Fig. 7d, e). The internal mould shows only inconspicuous riblets on the flanks. Some pieces of shell are preserved on the flank and venter, showing numerous fine lirae, crossing over the ventral area. The suture, partly preserved, shows a wide, deeply serrated external lobe and frilled saddles, the successive sutures overlapping on mid-flank ( Fig. 7f). Another fragment of a body chamber, from the same locality ( UJF. ID 10885), with Wh at about 45 mm, has preserved the shell, which is covered with fine lirae. These lirae are straight on the inner part of the flank, slightly concave adorally on the outer third, and crossing over the ventral area. The venter is relatively broad, slightly convex and bicarinated ( Fig. 7c). A larger fragment of body chamber from Viota ( UJF. ID 10884), measuring 110 mm of length, and preserved as an internal mould, shows an almost flat, bicarinate venter and inconspicuous lirae on the flanks ( Fig. 7a, b).

This material collected by BREISTROFFER (1952) seems to be very close to the unfigured specimen described as Platiknemiceras sp. A by ETAYO- SERNA (1979, p. 76, text-figs 8A, K). The poorly figured Platiknemiceras columbiana ETAYO- SERNA, 1979 seems to differ from both Platiknemiceras sp. A and the BREISTROFFER’ s material, only by its finely crenate ventrolateral edges.

The taxonomic position of those colombian ammonites remains unclear. Pending a revision of this group on the basis of new material, the morphological features of the suture line has led us to exclude these ammonites from either Platiknemiceras auctorum or Knemiceras .

Peru: CASEY (1961, p. 354, fig. 1c, d) published drawings of the ventral and cross-section views of a specimen attributed to Platiknemiceras , which was collected by HARRISON (1953, p.15) from the Early Albian of Canta, Peru (C.W. WRIGHT collection). We were unable to locate this specimen, which is supposed to be housed in the collections of the Natural History Museum of London.

ROBERT (2002, pl. 16, figs 3, 4) and ROBERT & BULOT (2004, pl. 2, fig. 4) have identified a specimen from the upper part of the Early Albian of Paria Puquio, Peru as Platiknemiceras cf. flexuosum KENNEDY, LANDMAN & COBBAN, 1998 . This poorly preserved ammonite, about 150 mm diameter, with a compressed whorl section, a narrow umbilicus, rounded and shallow umbilical wall, convex flanks, extremely narrow flat to slightly concave venter and lack of ornament, cannot be identified specifically, and may represent a primitive morphology of the genus Glottoceras HYATT, 1875 .

Japan: Platiknemiceras caseyi MATSUMOTO, 1980 (in MATSUMOTO, KANMERA & OHTA, p. 333, pl. 37, fig. 5), from the middle member of the Yatsushiro Formation, Kyushu, Japan, (?Early Albian) was described on the basis of a single specimen, 27 mm in diameter. It has a compressed whorl section, a narrow umbilicus and a bicarinate venter, and it is characterized by flexuous low ribs on the outer part of the flank, ending with ventrolateral clavi. This ribbing style is not characteristic of the genus Platiknemiceras , and it seems that this enigmatic specimen of inaccurate age does not belong to the Engonoceratoidea .