VOL. 31, N° 1, 2012 Revue De Paléobiologie, Genève (Juillet 2012) 31 (1): 159-170 ISSN 0253-6730

1661-5468

VOL. 31, N° 1, 2012 Revue de Paléobiologie, Genève (juillet 2012) 31 (1): 159-170 ISSN 0253-6730

Knemiceras gracile Douvillé, 1916: a misunderstood Early Albian ammonite from north Sinai (Egypt), and considerations on the genus Platiknemiceras Bataller, 1954

Jean-Louis Latil1 & Mohamed Fouad Aly2

Abstract The taxonomic treatment of the Engonoceratoidea Hyatt, 1900 (Wright et al., 1996) is problematic and confusing, and recently has gained the attention of many paleontologists (Robert, 2002; Bujtor, 2010; Bulot, 2010; Latil, 2011). It appears that most of the described genera within this superfamily were based on morphological features, without stratigraphic and/or phylogenetic control. In particular, the genus Platiknemiceras Bataller, 1954, seems to represent a classic example of this problem. Knemiceras gracile Douvillé, 1916, from the Early Albian of Maghara area, north Sinai, Egypt, referred by Casey (1961) to Platiknemiceras, appears to be an often misinterpreted species. Species identified as members of the genus Platiknemiceras Bataller, 1954 in the literature, from Iran, Lebanon, Egypt, Tunisia, Algeria, Spain, France Peru, Colombia, Venezuela, USA and Japan are discussed herein, in order to revise their taxonomic positions. The type species, Platiknemiceras bassei Bataller 1954, is redescribed. A re-examination of the relationships between Knemiceras gracile and other members of the genus Platiknemiceras strongly suggests that this latter genus is polyphyletic.

Keywords Albian, Ammonites, Engonoceratoidea, Platiknemiceras, Parengonoceras, Knemiceras, Egypt, Spain, Tethys.

I. Introduction for which their taxonomic positions are discussed below. Many ammonites with compressed whorl section and Knemiceras gracile Douvillé, 1916 was described from reduced ornament, have been subsequently attributed to the Albian of the Maghara area, north Sinai, Egypt from this morphological genus, including: Platiknemiceras a single specimen. Since that time, many specimens sequanense Destombes, 1979; Platiknemiceras have been reported in the literature as belonging to this colombiana Etayo-Serna, 1979; Platiknemiceras caseyi species. It has even been regarded as an index species for Matsumoto 1980; Platiknemiceras valencianum Mas a regional biozone of the Early Albian (Abu-Zied, 2008). & Wiedmann, 1980; and Platiknemiceras flexuosum Since Casey (1961), it has been placed within the genus Kennedy et al., 1998. The taxonomic positions of all Platiknemiceras Bataller, 1954, and it seems that this this material are herein reconsidered. taxon has been often misinterpreted and its application in Bulot (2010, p. 171) pointed out that the genus the literature is confusing. Platiknemiceras was most probably polyphyletic and The genus Platiknemiceras was originally created retained only the following species: Platiknemiceras by Bataller (1954) as a subgenus of Knemiceras bassei Bataller, 1954; Platiknemiceras hachourii for Knemiceras (Platiknemiceras) bassei Bataller, (Dubourdieu, 1953); and Platiknemiceras flexuosum 1954 (p. 175, fig. 1): A species that have a compressed Kennedy, Landman & Cobban, 1998. Bujtor (2010, section, narrow umbilicus and feeble, flexuous costation. p. 13) adopted an even more conservative conception of It was interpreted by Casey (1961) as a genus close to the genus, close to that of Casey (1961). Latil (2011, Parengonoceras Spath, 1924, but was characterized p. 349) regards the type species of Platiknemiceras as by a thinly discoidal shell, accompanied by an a direct ancestor of the genus Parengonoceras s.str. extreme reduction of sculpture. It should be noted that and placed it within this latter genus. Both Knemiceras Platiknemiceras bassei is known only from juvenile gracile and Platiknemiceras bassei are herein redescribed specimens, which are morphologically very close to the and discussed, in order to clarify the phylogenetic juvenile stages of growth of Parengonoceras species. relationships between the Albian species previously Casey referred several tethyan ammonites to this genus referred to the genus Platiknemiceras Bataller, 1954.

1 Le Maupas, F-05300 Lazer, France. E-mail: [email protected] 2 Geology Department, Faculty of Science, Cairo University, Giza, Egypt. E-mail: [email protected] 160 J.-L. Latil & M. F. Aly

II. Conventions Knemiceras gracile H. Douvillé, 1916 Figs 1, 2 All dimensions of specimens are given in millimetres: D = diameter, Wb = whorl breadth, Wh = whorl 1916. Knemiceras gracile H. Douvillé, p. 128, pl. 16, height, U = umbilical diameter, v = distance between fig. 9; text-fig. 42. the two ventrolateral edges. Figures in parentheses are ? 1940. Engonoceras gracile Douvillé H.- Basse, p. 438, pl. 4, fig. 1; text figs 9, 10. dimensions as a percentage of the diameter at the point 1956. Knemiceras gracile H. Douvillé, 1916.- of measurement. Mahmoud, text-fig.15. The following conventions are used to indicate the ? 1956. Knemiceras aff. gracile H. Douvillé.- Mahmoud, repositories of specimens mentioned in the text: p. 66, pl. 4, fig. 6. UJF.ID, Institut Dolomieu, Grenoble, France; FSL, non 1956. Knemiceras gracile H. Douvillé, 1916.- Paleontological collections of the Université Claude Arnould-Saget, p. 16, text-fig. 10; pl. 3, figs 3, 4. Bernard, Lyon, France; MHNP, Museum National 1961. Platiknemiceras gracile Douvillé.- Casey, p. d’Histoire Naturelle de Paris, France; MGSB, Museo 354. Geológico del Seminario de Barcelona, Spain; non 2006. Knemiceras gracile Douvillé, 1916.- Abu-Zied, fig. 2r, pl. 6, figs 10-12. NHML, Natural History Museum of London, England; non 2008. Knemiceras gracile Douvillé, 1916.- Abu-Zied, UNCMHN, Universidad Nacional de Colombia, Museo figs 2R, 7A-C. de Historia Natural, Bogota, Colombia. Holotype: The holotype by monotypy is the specimen figured by H. Douvillé (1916, pl. 9, text-fig. 42), from III. Systematic Paleontology NW of Talat el Fellahin (now Gabal Maaza), SE of Gabal Maghara, north Sinai, Egypt. The holotype, herein re- Order Ammonoidea Zittel, 1884 figured (Figs 1, 2), is deposited in the paleontological Suborder Ammonitina Hyatt, 1889 collections of the Université Claude-Bernard, Lyon as Superfamily Engonoceratoidea Hyatt, 1900 FSL.EM.1788 (former collections of the École des Mines Family Knemiceratidae Hyatt, 1903 de Paris). Genus Knemiceras Böhm, 1898 Dimensions of the holotype: [= Iranoknemiceras Collignon, 1981a, p. 258, type FSL.EM.1788: D: 53.0 (100) – Wh: 30.0 (57) – Wb: 16.0 species Knemiceras uhligi (Choffat) var. douvillei (30) – U: 5.0 (9) – v: 2.4 (5) – Wb/Wh: 0.53 Basse, 1940, p. 431, by original designation] FSL.EM.1788: D: 70.0 (100) –Wh: 38.0 (54) – Wb: 21.0 (30) – U: 6.5 (9) – v: 3.7 (5) – Wb/Wh: 0.55 Type species: Ammonites syriacus Buch, 1850, p. 20, Description: A single specimen preserved as calcareous by the original designation of Böhm, 1898, p. 200. internal mould of a subadult phragmocone, 70 mm in diameter, with an estimated total diameter of at least See full discussions in Bulot (2010) and Latil (2011). 140 mm. The discoidal shell shows involute coiling with

a b 1 cm c d

Fig. 1: The holotype of Knemiceras gracile H. Douvillé, 1916, FSL.EM.1788, NW of Talat el Fellahin (now Gabal Maaza), SE of Gabal Maghara, north Sinai, Egypt, Early Albian. Knemiceras gracile Douvillé, 1916: a misunderstood Early Albian ammonite (Egypt) 161

1 cm

Fig. 2: suture of the holotype of Knemiceras gracile 1 cm H. Douvillé, 1916, FSL.EM.1788, NW of Talat el Fellahin (now Gabal Maaza), SE of Gabal Maghara, north Sinai, Egypt, Early Albian (After Douvillé, 1926). a very narrow umbilicus, comprising 9% of the diameter. The umbilicus increases in diameter on the last stages of ontogenesis, as suggested by traces on the flanks of the last whorl of the phragmocone. The umbilical wall is shallow and vertical with sharp umbilical shoulder. The whorl section is compressed, subtriangular (Wb/ Wh between 0.53 and 0.55), with maximum width at about the inner third of the flanks. The flanks are gently convex, converging to a very narrow venter (ventral thickness comprising less than 5% of the diameter). The ventral area is grooved and smooth, limited by rows of Fig. 3: Knemiceras sp. ind., the specimen figured by Basse weak ventrolateral bullae, forming sharp, crenulated (1940, pl. 4, fig. 1; text fig. 9, 10), as Knemiceras ventrolateral edges. The lateral ornament is almost gracile, from unknown stratigraphical level of the nonexistent and restricted to weak, broad, coarse, Albian of Mdereidj, Lebanon. regularly alternating primary and intercalatory ribs, up to a diameter of 40-50 mm. The ribbing style, while not well expressed, is characteristic of the genus. Beyond 50 mm 16, text-fig. 10; pl. 3, figs 3, 4), from the Early Albian of in diameter, the lateral ribbing becomes inconspicuous, southern Tunisia, are too weathered to be fully identified. and tends to disappear altogether. The body chamber is Furthermore, the Tunisian specimens have a wider and unknown from the holotype. The suture shows entire flatter ventral area, and less convex flanks than the saddles and feebly indentated lobes (Fig. 2). holotype. Discussion: This species remains poorly known, but Abu-Zied (2006, fig. 2r, pl. 6, figs 10-12; 2008, figs seems to represent a primitive morphology of the genus 2R, 7A-C) mistakely figured a single specimen from Knemiceras Böhm, 1898, characterized by an almost El Tourkumanyia, north Sinai, Egypt, as Knemiceras lack of lateral ornament, a very narrowly grooved gracile. The figured specimen, showing strong ribbing, ventral area, and a suture with entire saddles and feebly wide umbilicus, pronounced umbilical tubercles, wide indentated lobes. flat venter, and frilled suture. These morphological The holotype seems to co-occur with an unequivocal features seem to fall within the variability of Knemiceras juvenile representative of the genus Douvilleiceras de spathi Mahmoud, 1956, rather than Knemiceras gracile Grossouvre, 1894 (Douvillé, 1916, p. 120; pl. 15, Douvillé, 1916. Therefore, the Knemiceras gracile fig. 7). Mahmoud (1956: p. 66) does not provide details biozone of Abu-Zied (2008) is not herein accepted and about the locality or stratigraphic level of the specimen needs to be revised. he figured, but Moret & Mahmoud (1953, p. 268 and Occurrence: Gabal Maaza, SE of the great Maghara 269) mentioned the species within their Levels 2 and 4, Dome, north Sinai, Egypt, Early Albian. This species indicating an Early Albian age. is doubtfully reported from the Albian of Mderedj, The specimens figured byB asse (1940, pl. 4, fig. 1; text- Lebanon. figs 9, 10), from an unknown stratigraphic level of the Albian of Mdereidj, Lebanon, seem to be close to the Genus Parengonoceras Spath, 1924 present species by their general shape, suture and narrow [= Platiknemiceras Bataller, 1954, type species concave ventral area, but the photographed specimen, Knemiceras (Platiknemiceras) bassei Bataller, 1954, herein figured (Fig. 3) is too weathered and corroded to p. 175, by original designation; = Platyknemiceras be specifically identified. Bataller, 1959, p. 65, nom. null.] The specimens figured by Arnould-Saget (1956, p. Type species: Amaltheus ebrayi Loriol, 1882, p. 7, by 162 J.-L. Latil & M. F. Aly

the original designation of Spath, 1924, p. 508. Robert (2002, p. 100) selected the specimen figured by Loriol (1882, pl. 1: MHNG.035492, Ebray collection) as lectotype (Pl. III, figs 1-4). The species was revised by

Latil (2008). 1 cm

See full discussions in Bulot (2010) and Latil (2011).

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. Fig. 4: The lectotype of Knemiceras (Platiknemiceras) bassei 1961. Platiknemiceras bassei Bataller.- Casey, p. 355, Bataller, 1954 (MGSB.11478), a reproduction of the fig. 1a-b. figure in Cazalda & Urquiola (1992, photo on page 1979. Knemiceras hachouri Masse & Thieuloy, p. 68. 75). ? 1982. Parengonoceras hachourii (Dubourdieu, 1953).- Renz, p. 31, pl. 3, fig. 4; text fig. 19c-d. 1992. Platiknemiceras bassei Bataller.- Cazalda & third of the flanks and tending to be projected outward Urquiola, p. 74, 1 fig. near the ventrolateral edge. The ribs, becoming fainter 1994. Platiknemiceras bassei Bataller.- Martinez, at the end of the preserved last whorl, are replaced by Grauges & Salas, pl. 3, fig. 4. growth striae. Both the body chamber and the adult 1996. Platiknemiceras bassei Bataller, 1954.- Wright, growth stages are unknown. The knemiceratid suture line Calloman & Howarth, p. 130, fig.100: 5a (not 5b). is diagnostic, with broad first lateral lobe, frilled lobes 1996. Platiknemiceras sp.- Wright, Calloman & and indentated saddles. This species seems to show a low Howarth, p. 130, fig.100: 5c. rate of growth, with short distance between two sutures, ? 2010. Platiknemiceras aff. bassei Bataller, 1954.- Bulot, p. 171, pl. 1, fig. 1. even on the internal whorls. Martinez et al. (1994, pl. 3, 2010. Platiknemiceras bassei Bataller, 1954.- Bujtor, p. fig. 4) figured a specimen from Les Ventoses, Marmellà, 13, fig.10, only the lateral view. Barcelona (MGSB 1044, probable paralectotype). This 2011. Parengonoceras bassei (Bataller, 1954).- Latil, p. juvenile specimen, 28 mm in diameter, shows rather 349, fig. 24; pl. 3, figs 37-41; pl. 4, figs 1-6. strong, prorsiradiate, sinuous riblets, rising on umbilical wall. Type material: The lectotype (MGSB.11478), An additional fragment, MGSB 8689, a 180° sector of a designated by Casey (1961, p. 23), has been figured by phragmocone, about 80 mm in diameter, is characteristic Bataller (1954, p. 175, figs 1, 2); Bataller (1959, p. of the species, with the ornament that tends to be effaced 65, fig. 826);C azalda & Urquiola (1992, p. 74, photo with age. The venter remains very narrow with age (v/D on page 75), herein figured (Fig. 4). = 0,045), becoming slightly convex. Dimensions of the lectotype: Discussion: This species represents the oldest known D: 60 (100) – Wh: 32 (53) – Wb: 17 (28) – U: 6 (10) – v: representatives of the genus Parengonoceras Spath, 3 (5) – Wb/Wh: 0.53 1924. It resembles superficially Knemiceras gracile Description: The species was described on the basis H. Douvillé, 1916 (p. 128, pl. 16, fig. 9), which has of two specimens identified from the Early Albian of ventrolateral bullae, coarse and wide ribs on the flanks of NE Spain (Tarragona). The lectotype is preserved as a the juvenile, and a suture with entire saddles. calcareous internal mould of a non-adult phragmocone. Knemiceras deserti Mahmoud, 1956 (p. 63, pl. 4, figs The coiling is involute, with a narrow umbilicus 3-5; text-figs 39-41) also shows a suture with entire comprising 10% of the diameter. The umbilical wall saddles. These two last species show strong affinities is shallow and almost vertical. The whorl section is with the genus Knemiceras s.str. Parengonoceras subtriangular, compressed, with a maximum width at the bassei is very similar in shape to Knemiceras hachourii umbilical seam. The flanks are feebly convex, converging Dubourdieu, 1953 (p. 23, pl. 2, fig. 10-18; pl. 3, fig. 1-5) to a narrow, bicarinate, slightly concave venter. Strongly, and differs mainly by its grooved ventral area, the lack of prorsiradiate, faint ribs arise to umbilical seam, tending true ventrolateral tubercles at a similar diameter. to blunt on the inner third of the flanks. The ribs curve Parengonoceras cf. hahourii (Dubourdieu) (Renz, backward at mid flanks, becoming radial on the outer 1982, p. 31, pl. 3, fig. 4; text-fig. 19 c-d) from the upper Knemiceras gracile Douvillé, 1916: a misunderstood Early Albian ammonite (Egypt) 163

part of Machiques Formation (? transition Aptian- Occurrence: Early Albian of Spain (Bataller, 1954), Albian), Quebrada Maraca, Venezuela, is characterized Tunisia (Latil, 2011) and maybe Venezuela (Renz, by a lack of lateral tuberculation, falcoid growth striae 1982). and bicarinate ventral area, and probably belongs to Parengonoceras bassei, differing only by its strongly frilled suture. IV. Discussion about The taxonomic The specimen recently figured by Bulot (2010, pl. 1, position of the taxa formely referred fig. 1) as Platiknemiceras aff. bassei Bataller, 1954, to the genus Platiknemiceras from the Aptian-Albian transition of Kuh-e-Bangestan Bataller, 1954 (Iran) is too poorly preserved to allow a determination. Bujtor (2010) re-figured two drawings of Casey Iran: Casey (1961, p. 354) reported three specimens (1961, fig. 1a, c) as “reproduction of the figure of Basse from the Albian of Hamiran, south Iran (Spath, 1931, (1954, p. 175)” (sic!). The lateral view corresponds to a p. 339, text-fig 111f-h; Casey, 1961, text-fig. 1c: drawing by Casey (1961, fig. 1a) of the original figure suture lines). Spath attributed these three specimens to of Bataller (1954, fig. 1), while the ventral view is a Knemiceras of the group of attenuatum-gabbi (sensu drawing of a peruvian specimen, identified by Casey lato). He interpretated them as probable new species, and (1961, fig. 1c) as Platiknemiceras sp. The same mistake considered them as passage-forms between Knemiceras is recorded in Wright et al. (1996, p. 130). and Engonoceras, following Pervinquière’s opinion Age: The type species Knemiceras (Platiknemiceras) (1907). These specimens are housed in the collections bassei Bataller, 1954 is known from Marmella, of the Natural History Museum of London under the Montmell Formation (Salou-Garraf, Tarragona). No other numbers C68402, C68403 and C648406 (Figs 5, 6). ammonites co-occur with this species in its type locality. These specimens, represented by rather large-sized Findings in the laterally equivalent Escucha Formation, phragmocones, show a compressed whorl section, narrow comprising “Hypacanthoplites milletianus (d’Orbigny), umbilicus, slightly convergent flanks and a narrow, Hypacanthoplites milletioides Casey and Sonneratia bicarinate venter. The ventrolateral edges are crenate to subglabra Casey” (Martinez, Grauges & Salas, tuberculate, and ventrolateral tubercles, when present, are 1994; Moreno-Bedmar et al., 2008), allow to assign an opposite on the venter. One specimen (NHML.C68406, Early Albian age to Platiknemiceras bassei (Leymeriella Fig. 6d-f) has radially elongated, periumbilical swellings, tardefurcata Zone to Sonneratia chalensis Zone). from which arise single, slightly flexuous, narrow, coarse Moreno-Bedmar et al. (2008) mention co-occurrence ribs that tend to be effaced with age. Morphological of ‘Hypacanthoplites’ and Parengonoceras bassei within features of this material strongly suggest close affinities the same bed in the basal Escucha Formation (Puntarro to the genus Knemiceras. Pending a revision of the south de Traiguera section, Cuenca del Maestrat, E Cordillera iranian engonoceratid faunas, and following Spath’s Ibèrica). This strongly suggests a basal Early Albian age opinion, we herein consider this material as belonging to for Parengonoceras bassei. one or several undescribed new species. 1 cm

a b c

Fig. 5: Knemiceras sp. nov., NHML.C68403, Albian of Hamiran, south Iran. 164 J.-L. Latil & M. F. Aly

a b c

1 cm

d e f

Fig. 6: Knemiceras sp. nov. a-c, NHML.C68402, Albian of Hamiran, south Iran; d-f, NHML.C68406, Albian of Hamiran, south Iran. Knemiceras gracile Douvillé, 1916: a misunderstood Early Albian ammonite (Egypt) 165

Lebanon: Knemiceras subcomplicatum Basse, 1940, p. specimen, from an unknown locality and stratigraphic 434, pl. 3, fig. 2, 3, from the Albian of Lebanon, is based level. This taxon is herein regarded as a nomen dubium. on two specimens. The holotype (MHNP.R052050: Geyer (1995, p. 8), placed Placenticeras saadensis pl. 3, fig. 2), from Bhamdoum, is a crushed, distorted Peron, 1890, p. 19; pl. 16, figs 3-7, within the genus and weathered fragment of an adult phragmocone, about Platiknemiceras. Breistroffer (1940a and b) regarded 130 mm diameter, with a compressed whorl section, this species as a Knemiceras. On the other hand, Latil narrow umbilicus and narrow venter. No ornament can (1989) identified it as an Engonoceras. This Late Albian be observed because of weathering. According to Basse, species from Algeria and Tunisia would be better placed the main features of this species are based mainly on the within the genus Hypengonoceras Spath, 1922, as a suture line, which is supposed to be characterized by deep result of its age and its sutural similarities. lobes, strongly frilled saddles and numerous accessory Tunisia: Casey (1961, p. 354), placed Knemiceras elements. The suture being strongly weathered, cannot hachourii Dubourdieu, 1953 (p. 23, text-fig. 7; be clearly studied. In addition, we agree with Young’s pl. 2, figs 10-18; pl. 3, figs 1-5) within the genus statement (1957, p. 2): “In my opinion, authors who Platiknemiceras. Latil (2011, p. 351) attributed this designate genera and/or species on sutures only are day- species to Parengonoceras. dreaming”. Spain: Platiknemiceras valencianum Mas & Wiedmann The paratype (MNHP.R52057, pl. 3, fig. 3), from (1980, p. 265, figs 6, 7A, 8A) from La Ermita de Chera Maaser-ech-Chouf, is a complete specimen, preserved section, Valencia Province, Spain (Late Albian), is based as a weathered, slightly distorted, calcareous internal on a poorly preserved fragment, but its suture shows strong mould, 115 mm diameter, comprising a 180° sector of affinities with sutures of Knemiceras sensu lato, such as body chamber. The whorl section is compressed, with a Knemiceras uhligi (Choffat, 1886). Furthermore, the maximum width at about mid flanks. The umbilicus is ventrolateral clavi of Platiknemiceras valencianum are narrow, with a shallow steep umbilical wall. The flanks alternating on the ventral area. The present authors are are slightly convex, the venter is narrow and bicarinate, satisfied to exclude this species from either Knemiceras tending to become convex and wider on the end of or Platiknemiceras auctorum. body chamber. The ornament on ventrolateral edges is France: Platiknemiceras sequanense Destombes, 1979, obscured, probably because of weathering. The inner part p. 114, pl. 4, fig. 3, was described from a single specimen of the last whorl (end of the phragmocone) shows feeble, from Quarry West, Bois du Perchois, Aube, France, coarse ribs arising by two or three from coarse umbilical level 1b (Early Albian, Sonneratia dutempleana zone, bullae. Basse (1940, p. 434) mentions numerous tiny, Cleoniceras floridum Subzone of Destombes). The ventrolateral tubercles on the phragmocone. holotype is a crushed and distorted fragment, represented Knemiceras subcomplicatum Basse, 1940, is referred by the end of the phragmocone and a 270° sector of to the genus Platiknemiceras by Casey (1961, p. 354) body chamber. The body chamber has been crushed without justification. From the main features of the before fossilisation. The umbilicus is very narrow, with paratype, this species should be placed within the rather deep, rounded umbilical wall. The whorl section genus Knemiceras, but we are not even sure that the is subtriangular, with maximum width just above the two specimens described by Basse belong to the same umbilical seam. Then the flanks are convergent to a species. The state of preservation of the holotype leads us narrow, slightly concave bicarinate venter (v/D = 0.05). to regard the present species as a nomen dubium. The author mentions smooth flanks, but an in-deep Egypt: Knemiceras deserti Mahmoud, 1956 (p. 63, examination of the type reveals the presence of faint, pl. 4, fig. 3-5, text fig. 39-41), was described from the flexuous growth striae on the body chamber. The suture Early Albian of Gabal Manzour, north Sinai, Egypt (see is poorly preserved, showing numerous, frilled elements. also Casey, 1961, p. 354), for small sized, compressed From the same level, Touch & Matrion (1995, pl. 3, ammonites, with subtriangular whorl section, narrow fig. 5) have figured an unregistered specimen identified umbilicus, steep umbilical wall, nearly smooth flanks, as Platiknemiceras sp. This incomplete specimen, 55 mm narrow venter, slightly crenate ventrolateral edges and in diameter, is characterized by a flat, narrow, bicarinate faint ventrolateral bullae that are opposite on venter. venter and strongly flexuous growth striae. Matrion This species is in need of a revision, but its suture (2010, p. 162, figs 116A-C) has figured two additional suggests strong affinities with the genus Knemiceras. A specimens from le Perchois quarries of Platiknemiceras careful examination of the paleontological collections sequanense. This species could be phylogenetically in Geology Department of Cairo University (Egypt), related to Parengonoceras bassei (Bataller, 1954). has convinced us that Knemiceras deserti represents USA: Platiknemiceras flexuosum Kennedy, Landman an extreme, compressed morphology of Knemiceras & Cobban, 1998, p. 38, fig. 51, 52 (Knemiceras sp. nov., douvillei Basse, 1940. Casey, 1961, p. 355), from the Glen Rose Limestone Algeria: Engonoceras jullieni Basse, 1940, p. 439, text- (Early Albian) of Texas, is characterized by its strongly fig. 11; pl. 4, fig. 4, from the Albian of Khenchela, Algeria, flexuous ribs, its convergent outer flanks and its narrow, is based on a poorly preserved, eroded and distorted slightly concave, bicarinate ventrer. This species is 166 J.-L. Latil & M. F. Aly

herein regarded as a primitive morphology of Texan of shell are preserved on the flank and venter, showing engonoceratoids and is provisionally placed within the numerous fine lirae, crossing over the ventral area. The genus Parengonoceras. However, we believe that the suture, partly preserved, shows a wide, deeply serrated north american engonoceratids most probably derive external lobe and frilled saddles, the successive sutures from some obscure Early Albian, tethyan stock, and overlapping on mid-flank (Fig. 7f). Another fragment of have evolved in isolation, on the Texan platforms. In our a body chamber, from the same locality (UJF.ID10885), opinion, this group deserves a separate generic status. It with Wh at about 45 mm, has preserved the shell, which should be noted that this species and Parengonoceras is covered with fine lirae. These lirae are straight on the roemeri (Cragin, 1893) are contemporaneous. inner part of the flank, slightly concave adorally on the Venezuela: Some discoidal specimens with a compressed outer third, and crossing over the ventral area. The venter whorl section and feeble ornament have been reported is relatively broad, slightly convex and bicarinated (Fig. from the Early Albian of Venezuela (Renz, 1982). 7c). A larger fragment of body chamber from Viota (UJF. Parengonoceras cf. hachourii (Dubourdieu) (Renz, ID10884), measuring 110 mm of length, and preserved as 1982, p. 31, pl. 3, fig. 4; text-fig. 19c, d: specimen an internal mould, shows an almost flat, bicarinate venter MB.Re2439), from the upper part of Machiques and inconspicuous lirae on the flanks (Fig. 7a, b). Formation in the Quebrada Maraca (Perijà river, This material collected by Breistroffer (1952) seems Venezuela), characterized by the lack of lateral to be very close to the unfigured specimen described as tuberculation, bicarinate ventral area, and prorsiradiate Platiknemiceras sp. A by Etayo-Serna (1979, p. 76, flexuous striae. It differs from Parengonoceras bassei text-figs 8A, K). The poorly figured Platiknemiceras mainly by its suture with strongly frilled saddles and columbiana Etayo-Serna, 1979 seems to differ from somewhat less convex flanks. This ammonite was both Platiknemiceras sp. A and the Breistroffer’s collected in the same stratigraphic levels of Prolyelliceras material, only by its finely crenate ventrolateral edges. gevreyi (Jacob, 1907), providing an Early Albian age, The taxonomic position of those colombian ammonites Leymeriella tardefurcata Zone to base of Douvilleiceras remains unclear. Pending a revision of this group on the mammillatum Superzone. basis of new material, the morphological features of the ?Parengonoceras sp. (Renz, 1982, p. 32, pl. 4, fig. 2), suture line has led us to exclude these ammonites from from the top of Machiques Formation, Quebrada either Platiknemiceras auctorum or Knemiceras. Macoita, Venezuela, is probably contemporaneous to Peru: Casey (1961, p. 354, fig. 1c, d) published drawings Prolyelliceras gevreyi (Jacob, 1907). This specimen, of the ventral and cross-section views of a specimen with compressed section, umbilical tubercles and attributed to Platiknemiceras, which was collected by flexuous striae, is close to Parengonoceras caneroti Harrison (1953, p.15) from the Early Albian of Canta, Collignon, 1981b, from which it differs by its strong Peru (C.W. Wright collection). We were unable to locate ventrolateral clavi that are opposite on the ventral area. this specimen, which is supposed to be housed in the Colombia: Platiknemiceras columbiana Etayo-Serna, collections of the Natural History Museum of London. 1979 from Cerro Bejucal, Colombia, basal part of the Robert (2002, pl. 16, figs 3, 4) and Robert & Bulot Capotes Member (?Middle Albian), is poorly figured (2004, pl. 2, fig. 4) have identified a specimen from the (p. 76, pl. 11, fig. 7: holotype by monotypy C-169/ upper part of the Early Albian of Paria Puquio, Peru as UNCMHNCPHt.N58). It is characterized by “A) finely Platiknemiceras cf. flexuosum Kennedy, Landman & crenate ventrolateral carinae, B) the trend of the bundles Cobban, 1998. This poorly preserved ammonite, about of lirae, radial on the central and adumbilical third of the 150 mm diameter, with a compressed whorl section, a flank, adorally concave on the adventral third” (Etayo- narrow umbilicus, rounded and shallow umbilical wall, Serna, 1979, p. 76). convex flanks, extremely narrow flat to slightly concave Breistroffer (1952, p. 2634) mentioned from the basal venter and lack of ornament, cannot be identified Middle Albian of Cundinamarca, Colombia, discoidal, specifically, and may represent a primitive morphology compressed and involute specimens attributed to the of the genus Glottoceras Hyatt, 1875. genus Knemiceras, with a costation reduced to very fine Japan: Platiknemiceras caseyi Matsumoto, 1980 (in striae and a narrow bicarinate venter. This material, placed Matsumoto, Kanmera & Ohta, p. 333, pl. 37, fig. 5), by Casey (1961, p. 355) in Platiknemiceras, has been from the middle member of the Yatsushiro Formation, borrowed by us from the collections of the Université Kyushu, Japan, (?Early Albian) was described on the Joseph Fourier (Grenoble, France). This material basis of a single specimen, 27 mm in diameter. It has consists of two fragments from Quipilé, and an additional a compressed whorl section, a narrow umbilicus and one from Viota, Quebrada Tasajera (Cundinamarca, a bicarinate venter, and it is characterized by flexuous Colombia). A small fragment of an internal mould of a low ribs on the outer part of the flank, ending with juvenile specimen from Quipilé (UJF.ID10886), about ventrolateral clavi. This ribbing style is not characteristic 30 mm long, shows a narrow, flat, bicarinate venter. It of the genus Platiknemiceras, and it seems that this has a compressed, subtriangular whorl section, with enigmatic specimen of inaccurate age does not belong to subparallel flanks (Fig. 7d, e). The internal mould shows the Engonoceratoidea. only inconspicuous riblets on the flanks. Some pieces Knemiceras gracile Douvillé, 1916: a misunderstood Early Albian ammonite (Egypt) 167

1 cm c

1 cm a b 1 cm

d e

1 cm

Fig. 7: ?Parengonoceras sp. nov. a, b, UJF.ID10884, Albian of Quebrada Tasajera, Viota, Cundinamarca, Colombia; c, UJF-ID.10885, Quipilé, Cundinamarca, Colombia; d-f, UJF-ID.10886, Quipilé, Cundinamarca, Colombia. 168 J.-L. Latil & M. F. Aly

V. Conclusion (J.-L.L.) to study the historical material from Egypt and Colombia; J.L. Dommergues (Dijon) and J.M. Pacaud A careful examination of the material previously (Paris) for allowing access to comparative material. We referred in the literature to the genus Platiknemiceras, are indebted to L. Percival (London), for providing to strongly suggests that this latter genus is polyphyletic. one of us (J.-L.L.), photographs of the iranian material Some species are herein placed within the genus housed in the NHML. We especially thank L.G. Bulot Parengonoceras Spath, 1924: (Marseille), B. Matrion (Troyes), K. Minor (Austin, - Knemiceras (Platiknemiceras) bassei Bataller, 1954 Texas) and J. Moreno-Bedmar (Barcelona) for useful (p.175, fig. 1). and friendly discussions, and K. Minor (Austin, Texas) - Knemiceras hachourii Dubourdieu, 1953 (p. 23, text- for improving the English manuscript. fig. 7; pl. 2, figs 10-18; pl. 3, figs 1-5). - Platiknemiceras sequanense Destombes, 1979 (p. 114, pl. 4, fig. 3). References Two species are referred to the genus Knemiceras Böhm, 1898: Abu-Zied, R.H. (2006) - Biostratigraphic framework of some - Knemiceras gracile H. Douvillé, 1916 (p. 128, pl. 16, Lower Cretaceous outcrops from northern Sinai, Egypt fig. 9; text-fig. 42). based on cephalopods. Egyptian Journal of Paleontology, - Knemiceras deserti Mahmoud, 1956 (p. 63, pl. 4, fig. 6: 125-158. 3-5, text fig. 39-41) Abu-Zied, R.H. (2008) - Lithostratigraphy and biostratigraphy One species seems to be closely related with the Late of some Lower Cretaceous outcrops from Northern Sinai, Egypt. Cretaceous Research, 29 (4): 603-624. Albian genus Hypengonoceras Spath, 1922: Arnould-Saget, S. (1956) - Contribution à l’étude des Engo- - Placenticeras saadensis Peron, 1890 (p. 19; pl. 16, noceratidae (les couches à Knemiceras du Sud-tunisien). figs 3-7). Annales des Mines Géologiques, 20: 47 p. Two species are regarded as nomen dubium Basse, E. (1940) - Les Céphalopodes des massifs côtiers - Knemiceras subcomplicatum Basse, 1940 (p. 434, pl. syriens. Notes et Mémoires du Haut-Commissariat de la 3, fig. 2, 3). République Française en Syrie et au Liban, 3: Etudes - Engonoceras jullieni Basse, 1940 (p. 439, text-fig. 11; paléontologiques, 411-472. pl. 4, fig. 4). Bataller, J.R. (1954) - Los Engonoceratidos en España. In: The taxonomic position of the following species has yet “Dr. D.F. Pardillo Vaquer homenaje postumo”, Facul- to be precised: tad de Ciencias de la Universidad de Barcelona: 173-178. Bataller, J.R. (1959) - Supplemento a las especies nuevas del - Platiknemiceras valencianum Mas & Wiedmann, Cretaceo. Cephalopoda. Boletin del Instituto Geológico y 1980 (p. 265, figs 6, 7A, 8A). Minero de España, 70: 65-66. ennedy andman - Platiknemiceras flexuosum K , L & Böhm, J. (1898) - Uber Ammonites pedernalis Von Buch. Zeits- Cobban, 1998 (p. 38, fig. 51, 52). chrift der Deutschen Geologischen Gesellschaft, 50: - Platiknemiceras columbiana Etayo-Serna, 1979 (p. 183-208. 76, pl. 11, fig. 7). Breistroffer, M. (1940a) - Révision des Ammonites du Vra- - Platiknemiceras caseyi Matsumoto, 1980 (in conien de Salazac (Gard) et considérations générales sur ce Matsumoto, Kanmera & Ohta, p. 333, pl. 37, fig. sous-étage albien. Travaux du Laboratoire de Géologie de 5). la Faculté des Sciences de l’Université de Grenoble, 22: Furthermore, the Iranian material reported by Spath 101 p. Breistroffer, M. (1940b) - Sur la découverte de Knemiceras (1931) and Casey (1961) as belonging to Knemiceras aff. saadense Thomas et Peron sp. dans le Vraconien de of group of attenuatum-gabbi (sensu lato) may represent Salazac (Gard). Comptes Rendus sommaires de la Société a new species of Knemiceras. The Peruvian material Géologique de France, 8: 87-89. identified by Robert (2002) and Robert & Bulot Breistroffer, M. (1952) - Sur la découverte de Knemicerati- (2004) as Platiknemiceras cf. flexuosum Kennedy, nae (ammonites albiennes) en Equateur, en Colombie et au Landman & Cobban, 1998, probably belongs to the Vénézuela. Comptes Rendus de l’Académie des Sciences, genus Glottoceras Hyatt, 1875 . 234: 2633-2635. Buch, L. von (1850) - Über Ceratiten. Physikalische Mathema- tische Abhandlungen der Koeniglichen Akademie der Wis- ACKNOWLEDGEMENTS senschaften zu Berlin, 1848: 1-33. Bujtor L. (2010) - Systematics, phylogeny and homeomorphy of the Engonoceratidae Hyatt, 1900 (Ammonoidea, Cre- The results presented here were obtained thanks to taceous) and revision of Engonoceras duboisi Latil, 1989. scientific agreements between the universities of Carnets de Géologie / Notebooks on Geology, Brest, Grenoble (France) and Cairo (Egypt) (Partenariat Hubert Article 2010/08 (CG2010_A08) Curien, Imhotep project 2009-10 n° 23167VJ and Bulot, L.G. (2010) - Appendix. Systematic palaeontology of STDF-IRD project 2011-2013). We thank A. Prieur Aptian and Albian ammonites from southwest Iran. In: (Lyon) and E. Robert (Grenoble), for allowing one of us Vincent, B., F.S.P. van Buchem, L.G. Bulot, A. Immen- Knemiceras gracile Douvillé, 1916: a misunderstood Early Albian ammonite (Egypt) 169

hauser, M. Caron, D. Baghbani & A.Y. Huc. Carbon- Contributions to Knowledge, n° 673. Washington D.C.: xi isotope stratigraphy, biostratigraphy and organic matter + 238 p. distribution in the Aptian – Lower Albian successions of Hyatt, A. (1900) - Cephalopoda. In: Zittel, K.A., Textbook of southwest Iran (Dariyan and Kazhdumi formations). Palaeontology, 1st English edition, translated by C.R. GeoArabia Special Publication, 4 (1): 167-195. Eastman: 502-592. Casey, R. (1961) - The Cretaceous (Albian) ammonite genus Hyatt, A. (1903) - Pseudoceratites of the Cretaceous. Mono- Platiknemiceras Bataller. Annals of the Magazine of graphs of the United States Geological Survey, 44: 352 p. Natural History, 13 (4): 353-357. Jacob, C. (1907) - Études paléontologique et stratigraphique Cazalda, S. & M.M. Urquiola (1992) - Catàlogo de los holo- sur la partie moyenne des terrains crétacés dans les Alpes tipos conservados en el Museo Geologico del seminario de françaises et les régions voisines. Thèse de la Faculté des Barcelona. Trabajos del Museo Geológico del Seminario, Sciences de Paris: 315 p. Barcelona, 223: 74-75. Kennedy, W.J., N.H. Landman & W.A. Cobban (1998) - Choffat, P. (1886) - Recueil d’études paléontologiques sur la Engonoceratid Ammonites from the Glen Rose Limestone, faune crétacique du Portugal. vol. I: Espèces nouvelles ou Walnut Clay, Goodland Limestone, and Comanche Peak peu connues. Travaux du Service Géologique du Portugal, Limestone (Albian) in Texas. American Museum Novi- 1: 1-40. tates, Museum of Natural History, 3221: 40 p. Collignon, M. (1981a) - Faune albo-cénomanienne de la For- Latil, J.L. (1989) - Les genres Engonoceras Neumayr and mation des marnes de Kazhdumi, région du Fars-Khuzes- Uhlig, 1881 et Hypengonoceras Spath, 1922 dans l’Al- tan (Iran). Documents du Laboratoire de Géologie de Lyon bien supérieur (z. à Dispar) de Salazac, Gard, France. Hors Série, 6: 251-291. Revue de Paléobiologie, Genève, 8 (1): 51-63. Collignon, M. (1981b) - In: Canérot, J. & M. Collignon. Latil, J.L. (2008) - A revision of Amaltheus ebrayi de Loriol, La faune albienne de Traiguera (province de Castellon- 1882, type species of the genus Parengonoceras Spath, Espagne). Documents du Laboratoire de Géologie de Lyon 1924. Revue de Paléobiologie, Genève, 27 (1): 249-264. Hors Série, 6: 227-249. Latil, J.-L. (2011) - Early Albian ammonites from Central Tunisia and adjacent areas of Algeria. Revue de Paléobio- Cragin, F.W. (1893) - A contribution to the Invertebrate paleontology of the Texas Cretaceous. Geological Survey logie, Genève, 30 (1): 321-429. of Texas, 4th annual report, 2: 139-294. Loriol, P. de (1882) - Études sur les faunes des couches du Gault de Cosne (Nièvre). Mémoire de la Société Paléonto- Destombes, P. (1979) - Les ammonites de l’Albien inférieur et logique Suisse, 9: 118 p. moyen dans le stratotype de l’Albien: gisements, paléonto- Mahmoud, G.I. (1956) - Études paléontologiques sur la faune logie, biozonation. In: Rat, P. et al. (Eds), L’Albien de crétacique du Massif du Moghara (Sinaï, Egypte). Publica- l’Aube. Les Stratotypes français, CNRS editions, 5: tion de l’Institut du désert d’Egypte, 8: 192 p. 51-193. Martinez, R., A. Grauges & R. Salas (1994) - Distribución Douvillé, H. (1916) - Les terrains secondaires dans le Massif de los ammonites del Cretácico inferior de la Cordillera du Moghara, à l’E de l’isthme de Suez. Mémoires de l’Aca- Costera Catalana e Ibérica Oriental. Cuadernos Geología démie des Sciences de Paris (2), 54: 1-184. Iberica, 18: 203-215. Dubourdieu, G. (1949) - Présence de Knemiceras compres- Mas, J.R. & J. Wiedmann (1980) - Ammoniten und Alter der sum Hyatt dans l’Aptien supérieur de l’Ouenza (Algérie). Mittelkreide-Transgression in der westlichen Pronvinz Bulletin de la Société Géologique de France, 5 (19): 26-27. Valencia, Spanien. Neues Jahrbuch für Geologie und Dubourdieu, G. (1953) - Ammonites nouvelles des Monts du Paläontologie Abhandlungen, 159 (2): 256-272. Mellègue. Bulletin du Service de la Carte Géologique Masse, J-P. & J.-P. Thieuloy (1979) - Précisions sur l’âge des d’Algérie, 1° série. Paléontologie, 16: 1-76. calcaires et des formations associées de l’Aptien sud- Dubourdieu, G. (1956) - Etude géologique de la région de constantinois (Algérie). Conséquences paléogéogra- l’Ouenza (Confins Algéro-Tunisiens).Publications du Ser- phiques. Bulletin de la Société Géologique de France, 7, vice de la Carte Géologique d’Algérie, 10: 659 p. 21 (1): 65-71. Etayo-Serna, F. (1979) - Zonation of the Cretaceous of Cen- Matrion, B. (2010) - Les ammonites. In: Colleté, C. (Coord.), tral Colombia by ammonites. Publicaciones Geológicas Stratotype Albien. Museum Nationale d’Histoire naturelle, Especiales del Ingeominas, 2: 1-188. Paris; Biotope, Mèze; BRGM, Orléans, 99-193. Geyer, O.F. (1995) - Knemiceras uhligi (Choffat) dans la Matsumoto, T. (1980) - In: Matsumoto, T., K. Kanmera & Chaîne Ibérique méridionale (Albien supérieur, Espagne). Y. Ohta. Cephalopod faunule from the Cretaceous Yat- Batalleria, 5: 5-17. sushiro Formation (Kyushu) and its implications. Transac- Grossouvre, A. de (1894) - Recherches sur la Craie supé- tions and Proceedings of the Palaeontological Society of rieure. Deuxième partie: Paléontologie. Les ammonites de Japan, N.S., 118: 325-338. la Craie supérieure. Mémoires du Service de la Carte Géo- Moreno-Bedmar, J.A., L. Bulot, J.-L. Latil, R. Martínez, logique Détaillée de la France, 1901: 264 p. O. Ferrer, T. Bover-Arnal & R. Salas (2008) - Preci- Harrison, J.V. (1953) - Geologia de la zona del camino entre siones sobre la edad de la base de la Fm. Escucha, mediante Canta y Huayllay. Boletin del Instituto de Geologia del ammonoideos, en la subcuenca de la Salzedella, Cuenca Peru, 9: 5-32. del Maestrat (E Cordillera Ibérica). Geo-Temas, 10: Hyatt, A. (1875) - The Jurassic and Cretaceous ammonites col- 1269-1272. lected in South America by Prof. James Orton, with an Moret, L. & G.I. Mahmoud (1953) - Nouvelles observations appendix upon the Cretaceous ammonites of Prof. Hartt’s stratigraphiques et paléontologiques sur l’Albien du massif collection. Proceedings of the Boston Society of Natural du Moghara (Sinaï-Egypte). Travaux du Laboratoire de History, 17: 365-372. Géologie de la Faculté des Sciences de Grenoble, 31: Hyatt, A. (1889) - Genesis of the Arietidae. Smithsonian 269-274. 170 J.-L. Latil & M. F. Aly

Péron, A. (1890) - Description des mollusques fossiles des ter- Spath, L.F. (1924) - On a new ammonite (Engonoceras iris sp. rains crétacés de la région sud des hauts plateaux de la n.) from the Gault of Folkestone. Annals of the Magazine Tunisie recueillis en 1885 et 1886 par M. Philippe Tho- of Natural History, 9(14): 504-508. mas. In: Exploration scientifique de Tunisie. Imprimerie Spath, L.F. (1931) - A monograph of the Ammonoidea of the Nationale, 1: I-XII, 1-103. Gault, part 8. Palaeontographical Society Monographs: Pervinquière, L. (1907) - Étude de paléontologie tunisienne. 313-378. 1: Céphalopodes des terrains secondaires. Mémoires de la Touch, R. & B. Matrion (1995) - Synthèse historique et géo- carte géologique de Tunisie: 438 p. logique de la carrière du Perchois-Ouest, Aube. Bulletin Renz, O. (1982) - The Cretaceous ammonites of Venezuela. annuel de l’Association Géologique auboise, 16: 15-24. Birkhaüser Verlag (Eds): 132 p. Wright, C.W., J.H. Calloman & M.K. Howarth (1996) - Robert, E. (2002) - La transgression albienne dans le Bassin Treatise on Invertebrate Paleontology. Part. L. Mollusca 4 Andin (Pérou): Biostratigraphie, Paléontologie (ammo- revised, Cephalopoda, Ammonoidea. Geological Society nites) et Stratigraphie séquentielle. Strata, 38: 380 p. of America and University of Kansas Press, 362 p. Robert, E. & L.G. Bulot (2004) - Origin, phylogeny, faunal Young, K.P. (1957) - Upper Albian (Cretaceous) Ammonoidea composition, and stratigraphical significance of the Albian from Texas. Journal of Paleontology, 31: 1-33. Engonoceratidae (pulchelliaceae, ammonitina) of Peru. Zittel, K.A. von (1884) - Cephalopoda. In: Zittel, K.A., Han- Journal of South American Earth Sciences, 17: 11-23. dbuch der Palaeontologie, Band 1, Abt. 2, Lief 3. Olden- Spath, L.F. (1922) - On Cretaceous ammonites from Angola, bourg. Munich & Leipzig, 329-522. collected by Prof. J.W. Gregory, D.Sc., FRS. Transac- tions of the Royal Society of Edinburgh, 53(1): 91-160. Accepté juin 2012