Contents
[Introduction] [Materials and methods]
[Historical overview ...]
[Systematic paleontology]
[Geographical and stratigraphical distribution ...]
[Mode of life ...]
[Phylogeny]
[Homoeomorphy]
[Conclusion]
[Appendix] [Bibliographic references] and ... [Table]
Department of Geology, University of Pécs, 6 Ifjúság Street, H-7624 Pécs (Hungary);
Corresponding address: 63 Fenyofa Street, H-2030 Érd (Hungary)
Manuscript online since December 24, 2010
The Engonoceratidae may well have originated on the shallow marine carbonate platforms of the southern margin of the Mediterranean Province of the Tethyan Realm during the
earliest Albian (?latest Aptian). In the entire Tethyan Realm and beyond adaptive radiation of the group was rapid and successful during the early Albian as indicated by endemic centres in the Peruvian Basin and the Western Interior Sea (USA). Later the group successfully enlarged its distribution, and invaded some provinces
(Mowry Sea, Canada) of the Boreal Realm. The mode of life of engonoceratids seems to have been
nektoplanktonic, epipelagic, and stenohaline, restricted to shallow water and platform or in some cases extremely shallow (littoral and
lagoonal) facies which may have helped their radiation. The group is distinctive and consists of nine genera and up to a hundred species, although its origin is still obscure. Their appearance may have been triggered by the oceanic anoxic event
(OAE 1b) and their rapid rise may have been helped by their shallow water mode of life and the global mid-Cretaceous warming and rise in sea level. The fall of the engonoceratids coincides with the
end-Cenomanian rapid transgression, which may have changed their shallow water habitats. Today the group is considered to have been a successful colonizer, a reliable stratigraphical indicator for shallow marine environments and a pioneer taxon for recognition of transgressive phases.
Based on new material and sutural analysis, Engonoceras duboisi is revised and assigned to Parengonoceras. A set of features consisting of a simplified ceratitid suture, lanceolate and compressed oxycone shell together comprising a shallow marine ecotype is pointed out as a successful and repeated morphotype among Mesozoic
Ammonoidea, for it was repeated three times during ammonoid evolution.
Ammonites; Albian; Cenomanian; phylogeny; systematics; homeomorphy.
L. (2010).- Systematics, phylogeny and homeomorphy of the Engonoceratidae , 1900 (Ammonoidea, Cretaceous) and revision of Engonoceras duboisi , 1989.- Carnets de Géologie / Notebooks on Geology, Brest, Article 2010/08 (CG2010_A08)
Systématique, phylogénie et homéomorphie
des Engonoceratidae , 1900
(Ammonoidea, Crétacé) et révision de Engonoceras duboisi
, 1989.- Les Enganoceratidae apparaissent à l'Albien basal (ou peut-être dès l'Aptien terminal) dans les séries de
plates-formes carbonatées de la marge sud de la province méditerranéenne du domaine téthysien. Leur efficacité évolutive leur permet de coloniser rapidement l'ensemble du domaine téthysien et même au delà comme le montre la présence de groupes endémiques dans le Bassin Péruvien et dans la mer intérieure occidentale des États-Unis. Plus tard, le groupe élargira encore plus son aire de distribution en envahissant certaines régions du domaine boréal (mer de Mowry, Canada).
Les Enganoceratidae semblent avoir eu un mode de vie nectoplanctonique, épipélagique et sténohalin au sein de mers chaudes peu à très peu profondes (rivages, lagons), dont la fréquence a favorisé leur évolution. Ce groupe, dont l'origine est obscure, est bien individualisé. Il comprend neuf genres et une centaine d'espèces. Leur apparition semble avoir été provoquée par la crise anoxique OAE 1b et leur rapide expansion favorisée par l'extension progressive des mers chaudes et épicontinentales qui accompagne la montée des eaux du milieu du Crétacé. Le déclin des Enganoceratidae semble lié à l'accélération de la montée des mers qui caractérise la fin du Cénomanien, sans doute par modification substantielle de leurs habitats peu profonds. Ce groupe est considéré comme un colonisateur efficace et un bon indicateur de milieux de faible profondeur en ambiance transgressive.
À partir de nouvelles études stratigraphiques et structurales, il est proposé une révision de l'espèce
Engonoceras duboisi qui doit être rattachée au genre Parengonoceras. Par ailleurs, il est mis en évidence que le fait de présenter des sutures simplifiées de
type Cératite et une coquille lancéolée à oxycône comprimé est symptomatique
d'ammonites adaptées à des milieux marins peu profonds. Ce modèle est, en effet, répété trois fois au long de l'évolution des Ammonoïdes
mésozoïques.
Ammonites ; Albien ; Cénomanien ; phylogénie ; systématique ; homéomorphie.
Engonoceratid ammonites form a distinct but perplexing group among Cretaceous Ammonoidea. Their systematics is still confusing and their origin is obscure. After the period of great monographs on engonoceratid ammonites (, 1903; , 1907; , 1910; , 1940; in et alii, 1947; , 1953; , 1955; , 1956; , 1970, 1982; , 1979), twelve new species have been introduced and forty species invalidated. This prompts a review of the group, and a discussion on the ecology and facies dependency of certain of its taxa. Recent debates on the systematics of Engonoceratidae also invite a review of the systematic position, ecology, distribution, phylogeny and homoeomorphy of this group of fossils and of the species assigned it.
The following abbreviations indicate the repositories of specimens mentioned in the text: HNHM, Hungarian Natural History Museum, Budapest, Hungary; LC, Lumachella Collection (private collection of Mr. Zoltán , 29 Szatmár Street, Budapest, H-1194 Hungary); MHNG-GEPI, Collection Muséum d'Histoire naturelle de Genève, Switzerland. Suture terminology is after & (1970): E, external lobe; L, lateral lobe; U, umbilical lobe; I, internal lobe. Note that et alii (2003) revised the sutural terminology of the Paleozoic prolecanitid ammonites, which implies its modification for Mesozoic ammonoids too, but this modification has not been published yet. All dimensions are in millimetres: D, diameter; Wb, whorl breadth; Wh, whorl height; U, umbilicus; myr: million year. Species names in brackets indicate invalidation.
The name Ceratites was introduced by de (1825) for a group of Triassic ammonites with simplified suture lines. As early research developed, Ceratites-like fossils of Cretaceous age were discovered, so early palaeontologists included these fossils into the ceratitid group, naming them as Cretaceous Ceratites (, 1848, p. 30) or "pseudo-ceratites based on an apparent resemblance between the sutures of Triassic ceratitids and those of Cretaceous engonoceratids" (, 1903; G. , 1940a, 1940b; & , 1976; , 1981; et alii, 1998a). The first engonoceratid ammonite was published by d' (1841) as Ammonites Vibrayeanus [= Neolobites vibrayeanus] from France, and by (1848) as Ammonites Syriacus [= Knemiceras syriacum] from Lebanon, and Ammonites pierdenalis [= Engonoceras pierdenalis] from Texas. (1848) also called attention to the importance of the group referring to them as Cretaceous ceratitids ('Kreide-Ceratit'). Although (1848) reported other "ceratitids" (whose suture lines superficially resemble those of the Triassic Ceratites) such as Ammonites Ewaldi and Ammonites Robini [= Metatissotia ewaldi and M. robini], these species belong to discrete taxa (Acanthoceratoidea) with no direct phyletic connection to the engonoceratids. The family name Engonoceratidae was introduced by (1900, p. 585) to include Protengonoceras, Engonoceras, Metengonoceras, and Neolobites. Later (1903, p. 144) established the family Knemiceratidae for Knemiceras. Based on the suture line (1912, 1928, 1931) placed the group in the Pulchelliidae, an assignment accepted by G. (1940b, p. 1066). (1922, 1924) introduced Hypengonoceras, and Parengonoceras. The last described member of the family is Platiknemiceras (, 1954).
In the mid 1900's (1941) and (1942) referred certain placenticeratid ammonites (Placenticeras mintoi, and P. planum) to the Engonoceratidae and (1956, p. 486) regarded Neolobites as incertae sedis again an indication of taxonomic uncertainty. Hypengonoceras was regarded as a placenticeratid ammonite (, 1960) at that time the only known Lower Cretaceous placenticeratid. Later (1970) summarized the systematic debates and content of the family, and & (1989, p. 365) revised the systematic position of the most discussed member, Hypengonoceras, which they regarded as having been derived probably from Parengonoceras. et alii (1996, p. 126) did not accept this rationale proposing a different origin for the genus: "Hypengonoceras, despite some resemblances to Engonoceratidae, seems to belong here" [= Placenticeratidae , 1900] and confirmed that the family had an uncertain origin, "but some genera closely resemble Pulchelliidae but are probably not closely related" ( et alii, 1996, p. 130) and placed the family in the Hoplitoidea with these eight genera included: (Engonoceras, Hypengonoceras, Knemiceras, Metengonoceras, Neolobites, Parengonoceras, Platiknemiceras, and Protengonoceras). At present, the systematic position of Hypengonoceras is again disputed ( & , 2004). (2002) resurrected the genus Glottoceras 1875, a view supported by & (2004, p. 20). As yet there is no unequivocal approach regarding the systematic position of the group. (1968, p. 747) had already emphasized the impossibility of placing the group properly until its sutural development is clearly known. He refers this family doubtfully to the Pulchelliidae or the Hoplitidae. (1970) gave a detailed analysis of the engonoceratid sutural history. In the past forty years many species were introduced but no significant work was done on the sutural ontogeny of the Engonoceratidae. Although & (2004) discussed the systematic position of the group placing it in the Pulchelliatoidea, this attribution is not yet widely accepted. et alii (2004), (2004), & (2005), et alii (2007), and et alii (2008) place the group in the Hoplitoidea, while (2008) refers it to the Pulchelliatoidea. Most recently, (2010, p. 169) rejects the pulchelliatoid origin of Engonoceratidae, suggesting that the genus Subpulchellia/Mogharaeceras is most likely an endemic offshoot of Barremites so cannot be the ancestor of the Engonoceratidae. Further uncertainty is indicated by (in , 2008b, p. 160) regarding the systematic position of European Engonoceras sensu stricto. & (2004, p. 11) stressed a pulchelliatoid origin based on a probable relation with the Barremian/Aptian Subpulchellia, but did not include a thorough discussion. (2010) proposes a new taxonomic rank for engonoceratid ammonites, the Superfamily Engonoceratoidea , 1900. This taxonomic approach has certain advantages: on one hand it emphasizes the integrity of the group, thus highlighting its morphological independence. On the other hand this approach leaves open the problem of the origin of the group. As mentioned, (2010) rejected the pulchelliatoid origin of engonoceratid ammonites. If so, the most plausible explanation for its origin is the hoplitoids. This view may be strengthened by new data on the ecology of Aptian Parahoplites ( et alii, 2009). Consequently, the systematics and origin of Engonoceratidae are still open issues. Recently Engonoceratidae was considered to consist of nine genera (Engonoceras, Glottoceras, Hypengonoceras, Knemiceras, Metengonoceras, Neolobites, Parengonoceras, Platiknemiceras, and Protengonoceras), and 143 species (Table 1) of which 38 are invalid. They are listed in the section discussing the described genera and their species. The genus Pseudengonoceras referred to by (1981, p. 517) is a typing error for Parengonoceras (P. , pers. comm., 2009).
Order Ammonitida , 1884
Suborder Ammonitina , 1889
Superfamily Hoplitoidea , 1890
Family Engonoceratidae , 1900
= Knemiceratidae , 1903, p. 144; Neolobitinae & 1958, p. 125
Genus Glottoceras , 1875, p. 372
(Fig. 1 
)
Type species. Glottoceras attenuatum , 1875, from the Lower Albian of Peru.
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Figure 1: Glottoceras attenuatum (, 1903). Reproduction of the original specimen of (1903, pl. 17, figs. 13-14).
Diagnosis. Typical engonoceratid coiling with narrow umbilicus. Cross section is rather compressed and pentagonal, or oval in some species. Venter is flat to slightly concave or rounded in adult stage. Ornamentation consists of rather strong, sometimes coarse prorsiradiate ribs, which may originate at the umbilical shoulder from bullae. Ventrolaterally bullae may also appear. Suture consists of many adventive and auxiliary elements. Relative breadth of saddles and lobes are similar. Saddles are bifid, lobes are finely frilled.
Occurrence. Glottoceras (sensu , 2002) is restricted to the Albian of South America.
Included species:
Remarks. (1956, p. 449) had already pointed out that South American 'Knemiceras' are different from the Knemiceras attenuatum described by (1940). On the basis of Art. 23.9 of (1999), (2002) resurrected Glottoceras to group the South American species of Knemiceras. (2002) and & (2004, p. 20) consider the genus Glottoceras to be restricted to these Andean species. In the assignment of species to this group, Knemiceras andinum must be included in Glottoceras if the rationale of & (2004, p. 20) is accepted.
Genus Engonoceras & ,
1881, p. 140
= Engonhoplitoides , 1940, p. 441; Epigonoceras ,
1956, p. 400
(Fig. 2 )
Type species. Ammonites pierdenalis , 1848, from the Cretaceous of Fredericksburg, Texas.
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Figure 2: Engonoceras pierdenalis (, 1848). Reproduction of the original figure of (1848, p. 31, pl. 6, figs. 8-10).
Diagnosis. Diagnosis is after et alii (1996, p. 130): "Involute, inner whorls very compressed with narrow, flat or sulcate venter; later flexuous striae normally repeated by weak, flat, flexuous or straight ribs ending in small ventrolateral clavi placed alternately and, in some shells, joined across venter by zigzagging ribs; umbilical and lateral tubercles may be present, and venter of last whorl may be rounded. Suture with more elements than Knemiceras and with saddles normally all entire except that outermost are bifid; external lobe normally with strongly divergent branches."
Occurrence. Engonoceras is widespread in its distribution. It is abundant with many species in North America (Colorado, Kansas, New Mexico, Oregon, Texas) and in Mexico (Sonora) and South America (Colombia, Peru) but also in the Tethys: North Africa (Algeria, Morocco, Tunisia), the Near East (Lebanon, Syria) and Asia (Borneo). It is also reported from Europe (England). Stratigraphically it ranges from the Early Albian to the Middle Cenomanian.
Included species:
Remarks. Although (1848, pl. 6, fig. 10) in his description of the type species indicates that all the saddles of the suture are bifid: "the saddle is rounded, however with one small secondary lobe in the middle", subsequent designations refer to entire or nearly entire saddles with only one bifid element being typical for Engonoceras ( et alii, 1996; et alii, 1998a, p. 5). According to (1956, p. 444) most species of Engonoceras have sutures with entire saddles and are ornamented with three or two rows of tubercles. In some species (e.g. E. complicatum) there are more (up to five) bifid or sometimes trifid saddles (E. elegans). Saddles are generally rounded, lobal necks are narrow. Saddles are always broader than lobes. (1940, p. 441) established Engonhoplitoides on the basis of the subdivision of the lateral lobe of the suture line, remarking that the shell morphology of Engonhoplitoides is closer to that of Engonoceras, while its suture line is more nearly like that of Knemiceras. This separation was discredited by et alii (1996).
Genus Neolobites , 1882, p. 389
(Fig. 3 )
Type species. Ammonites Vibrayeanus d', 1841, p. 322, from the Cretaceous of Sarthe (France).
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Figure 3: Neolobites vibrayeanus (d', 1841). Reproduction of the original figure of d' (1841, p. 322, pl. 96, figs. 1-3).
Diagnosis. Highly variable, oxycone and lanceolate engonoceratid, with small, shallow umbilicus. Suture is extremely simplified consisting of rounded, narrow lobes and wide rounded saddles. Cross section is variable from compressed to slightly inflated. Venter is variable from wide to narrow trapezoidal or simply rounded and in some species ornamented by fine crenulations. Sculpture is variable, too, ranging from smooth, unornamented forms to flexuous and ventrally ornamented forms.
Occurrence. Neolobites is restricted to the Middle (N. fourtaui) and Late (N. vibrayeanus) Cenomanian being a successful shallow marine taxon among Engonoceratidae. The genus is characterized by distinct morphotypes and populations in the Tethyan Realm and seamingly preferred lower latitudes and subtropical/tropical settings ( & , 2005, p. 942). It is reported from South America (Bolivia, Colombia, Peru, and Venezuela), Africa (Algeria, Egypt, Libya, Morocco, Niger, and Tunisia), Europe (France, Portugal, and Spain), and the Near and Middle East (Israel, Jordan, Lebanon, Oman, and Syria).
Included species:
Remarks. & (2005) stated that only N. vibrayeanus, N. fourtaui and N. peroni deserve discrete specific status. et alii (1981, p. 28) convincingly ruled out N. bedoti from Neolobites. It was placed in Metengonoceras ( et alii, 1992). Reported new species are based on fragmented and worn specimens. Therefore the sculpture and suture appear to be simplified (e.g. N. bassleri, N. kummeli). & (1981, p. 24) argued that N. peroni is also no more than an intraspecific variety of N. vibrayeanus. and (2005) did not discuss the earlier authors' opinion, but maintain N. peroni as a distinct species based on slight morphological differences between N. vibrayeanus and N. peroni. Taking into consideration the extreme morphological variety of the populations of Neolobites, the present author accepts the opinion of & (1981) on N. peroni, and considers this species an intraspecific variety of N. vibrayeanus. It is possible that N. fourtaui will also be found to fall into the intraspecific range of N. vibrayeanus, especially taking into consideration the eco-morphological analysis of & (2005). The other possibility, that future research will elucidate, is that N. fourtaui and N. vibrayeanus form two discrete chronospecies, namely N. fourtaui is older and N. vibrayeanus is younger. The Early Cenomanian N. bedoti of (1912, p. 31) has been assigned to Metengonoceras by & (1984, p. 105) because of the very similar morphologies of Neolobites and Metengonoceras. Based on these similarities, the most probable ancestor of Neolobites is Metengonoceras ( & , 2001).
Genus Knemiceras ,
1898, p. 200
= Cnemidoceras , 1900, p. 24; Cnemioceras ,
1900, p. 85; Knemoceras ,
1902, p. 7;
Omaimaiceras , 1955; Iranoknemiceras ,
1981, p.
258
Type species. Ammonites Syriacus , 1848, from the Neocomian of Lebanon
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Figure 4: Knemiceras syriacum (, 1848). Reproduction of the original figure of (1848, p. 21, pl. 6, figs. 1-3).
Diagnosis. Diagnosis is after et alii (1996, p. 130): "compressed to moderately inflated; sides flat and parallel or converging; venter flat or slightly concave; ribs moderately to very strong, sparse, rounded or flat, arising singly or in pairs from stout umbilical tubercles; ending in ventrolateral clavi or crossing venter. Suture with frilled lobes and slightly frilled, rarely entire saddles; commonly irregular."
Click on thumbnail to enlarge the image.
Figure 5: Highly variable cross sections of Knemiceras species. A.) K. compressum (, 1903, pl. 16, fig. 16); B.) K. compressum (, 1970, text-fig. 9c); C.) K. subcompressum ( et alii, 1997, fig. 2f); D.) K. kazhdumiense (, 1981, pl. 3, fig. 1c); E.) K. douvillei (, 1981, pl. 4, fig. 1b); F.) K. persicum (, 1981, pl. 5, fig. 1b).
Occurrence. Separation of South American forms of Knemiceras as Glottoceras by (2002) does not restrict the geographic distribution of the remaining Knemiceras species, which are reported from Europe (France, Portugal, Spain), North Africa (Algeria, Egypt, Tunisia), Near East (Israel, Lebanon, Syria), Middle East (Iraq, Iran, Oman), and also from South America (Colombia, Peru). Stratigraphically, Knemiceras appears in the earliest Albian (?latest Aptian) and disappears in the Early Cenomanian.
Included species:
Remarks. Resurrection of Glottoceras by (2002) did not solve the systematic uncertainties around Knemiceras. He considered the genus to be a geographical variant and leaves open the problem of the systematic position of the remaining Knemiceras species (e.g. K. attenuatum spinosum from Australia: , 1963). Other occurrences of 'Knemiceras' attenuatum from Tunisia (, 1956, pl. 2, fig. 3a-b) and Iran (, 1981, pl. 6, fig. 2) should be convincingly placed somewhere in the Engonoceratidae. & (1989, p. 364) proposed that Hypengonoceras ibericum be placed in Knemiceras on the basis that quadrituberculate ornament is unknown in Hypengonoceras. Bifid (e.g. K. compressum, K. subcompressum) or irregular (K. douvillei, K. iraniense) saddles are present in the suture line at any growth stage, distinguishing all Knemiceras species from Engonoceras and Platiknemiceras. The extreme intraspecific variation seen in Knemiceras ( & , 1989, p. 383) especially in its inflation and ornamentation ( & , 1991) is easily observable in the cross- sections of the body chambers of various species (Fig. 5). Cross-sections of Knemiceras range from the slender lanceolate forms (K. compressum) through wider (K. kazhdumiense, K. subcompressum) and subcircular (K. douvillei) and rectangular (K. persicum) forms. (1961) decreased this great variation within the genus by distinguishing the thin, unsculptured and less ornamented forms as Platiknemiceras. The extreme variability of shell morphology of Knemiceras may have been linked to the mode of life.
Genus Protengonoceras , 1903, p. 153
(Fig.
6 )
Type species. Engonoceras Gabbi , 1898, p. 197, from the Cretaceous of Texas (USA).
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Figure 6: Protengonoceras gabbi (, 1898). Reproduction of the original specimen of (1898, p. 197).
Diagnosis. Diagnosis is after et alii (1996) and et alii (1998a). Conch is like Engonoceras but lacks tubercles at any stage and has feeble, smooth ribs only on adult stage. Cross section is compressed to very compressed and tabulate or may be rounded at adult stage. Where ventrolateral shoulder is present, it is always sharp. The suture line is almost straight and consists of many low adventive saddles, which are entire. Adventicious lobes are narrow and incised.
Occurrence. Protengonoceras is reported from the Early and Middle Albian of Peru, Texas, Mexico and Egypt. (1981) reported two species from the latest Albian and earliest Cenomanian of Iran.
Included species:
Remarks. (1903, p. 157) referred 's Sphenodiscus emarginatus to both Protengonoceras and Engonoceras. Without detailed description it can only be considered a nomen nudum so cannot be maintained. (1903, p. 157) referred to this ambiguity: "not having seen any specimens of the species, I can not say positively that it is a member of this genus", therefore this reference is omitted. (1981) reported two new species from the Late Albian-Early Cenomanian of Iran. However, all other species are reported from the Early and Middle Albian, and no occurrences are known from the Late Albian or younger. (1981) introduced P. prestati possessing well preserved dorsolateral and lateral bullae and a suture line with bifid saddles typical of Knemiceras. Protengonoceras lacks tubercles and never presents bifid and/or incised saddles. On that basis, P. prestati belongs rather to Knemiceras; however the single figured specimen (, 1981, pl. 6, fig. 3a-b) does not permit a reliable revision.
Genus Metengonoceras ,
1903, p. 179
= Epengonoceras , 1924, p. 508
(Fig. 7
)
Type species. Metengonoceras inscriptum , 1903, from the Albian of Decatur, Texas.
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Figure 7: Metengonoceras inscriptum , 1903. Reproduction of the original figure of (1903: p. 180, pl. 25, figs. 5-9).
Diagnosis. Diagnosis is after (1987, p. C2). Metengonoceras includes very compressed, almost smooth ammonites having very narrow umbilicus and narrow, flattened venters, which may be rounded on the adult body chamber. Sculpture consists of falcoid growth lines. On some species, weak arcuate ribs appear a little above the midflank. Rarely faint umbilical bullae may appear. Suture is simplified, consisting of closely shaped numerous auxiliary and adventive elements. Most of the saddles are wide, being undivided and rounded. Larger lobes may be frilled moderately.
Occurrence. Metengonoceras occurred in the Gulf of Mexico during the Middle Albian ( & , 1989). Many species flourished in the Western Interior of the USA. Repeated speciation took place there (R.W. , 2007) from which certain Metengonoceras species invaded northward (Mowry Sea, Canada) and eastward (Normandy, France). During the Cenomanian Metengonoceras enlarged its territory outside North America. It is reported from Europe (France, Germany), Africa (Algeria, Egypt, Niger, Nigeria, Tunisia), and the Middle East (Jordan) as well as from South America (Brazil, Ecuador) reached through the Saharan seaway. Metengonoceras tended to become more acute and sharply keeled during the Cenomanian.
Included species:
Remarks: (1952) extensively reviewed the type species of Epengonoceras (E. dumbli) and concluded that the genus is not separable from Metengonoceras. This view was accepted by & (1984, p. 100) and is also followed here. R.W. (2007) reports stratigraphical ranges of engonoceratid ammonites from North America showing that certain Metengonoceras species occur in the Late Albian (M. teigenense, M. aspenanum) and Cenomanian (M. dumbli, M. acutum) suggesting that in North America Metengonoceras species range from earliest Late Albian to the latest Cenomanian.
Genus ? Hypengonoceras , 1922, p. 112
(Fig. 8 )
Type species. Placenticeras Warthi , 1895, from the Albian Utatur Group of southern India.
 |
 |
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Figure 8: Hypengonoceras decaryi (, 1963). Reproduction of the figures of & (1989: figs. 112-113).
Diagnosis. Diagnosis is after & (1989, p. 362). "Umbilicus is narrow. The shell compressed, the flanks generally little rounded. Sculpture consists of low falcoid ribs, often very weak and only visible under oblique lighting. Umbilical tubercles may be present, but are never very prominent. Venter is flat to concave with alternating ventral clavi at least at some stage. Dimorphism pronounced, but mainly restricted to differences in size. Suture variable; in typical forms some saddles are little divided and 'pincer-like' with bifid folioles; in others as incised as in a normal placenticeratid pattern."
Occurrence. Hypengonoceras is reported from the Late Albian of Europe (France), Africa (Madagascar, Morocco, Mozambique, Zululand), Near East (Israel), India, and probably Sakhalin (Japan). & (1984, Table 1) noted a doubtful occurrence in the Late Cenomanian of the Utatur Group, South India but they did not figure or describe it, so this occurrence is omitted.
Included species:
Remarks. Although & (1989) indicated the placenticeratid-like pattern of the suture of Hypengonoceras, they assigned the genus to the Engonoceratidae. This clearly relates to the debate on the systematic position of Hypengonoceras for it is the most disputed among the Engonoceratidae and is again under reconsideration ( & , 2004). This continuing dispute is based on the general differences between the Engonoceratidae and the Placenticeratidae. The Engonoceratidae are characterized by simplified ceratitic saddles (best seen on all species of Protengonoceras, Engonoceras, Metengonoceras, and Neolobites) in contrast to the frilled and finely incised saddles and lobes of Placenticeratidae. In this scheme, Parengonoceras and Hypengonoceras are transitional between the Engonoceratidae and the Placenticeratidae ( & , 1989, p. 364); however the first-named genus was placed in the Engonoceratidae without dispute - partly because of its known stratigraphical distribution (Early and Middle Albian) and because Parengonoceras possesses two true adventive lobes (, 1970). Only H. ibericum is questioned as a member of the genus by & (1989, p. 364) because quadrituberculate ornament is unknown in Hypengonoceras. They proposed that H. ibericum be placed in Knemiceras. Since the thorough discussion of Hypengonoceras by & (1989) no significant progress toward clarifying the position of the genus has been published; therefore the opinion of & (1989) is maintained here.
Genus Parengonoceras , 1924, p. 508
(Fig. 9 )
Type species. Amaltheus Ebrayi , 1882, from the Albian of Cosne, Nièvre, France.
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Figure 9: Parengonoceras ebrayi (, 1882). Reproduction of the original figure of (1882, p. 7, pl. 1).
Diagnosis. Coiling is typically engonoceratid. Conch is evolute. At early stages the cross section is lanceolate with tabulate venter. The lanceolate cross-section varies between wide triangular to acute and compressed. At adult stage the cross-section remains lanceolate or becomes oval. Venter is smooth or slightly grooved. Umbilical shoulder is low and always rounded and unornamented. Flanks are smooth in most cases or ornamented by ventrolateral, lateral or dorsolateral bullae. Falcoid ribs may appear ventrolaterally. Venter is usually narrow or in some species wide at adult stage and ornamented by weak tubercles. In some species these tubercles develop strongly. Suture line consists of two adventive and many auxiliary elements. Lobes are always irregularly incised. Saddles are never entire, always weakly or strongly incised.
Occurrence. Although Parengonoceras is widely considered to be restricted to the Early and Middle Albian ( & , 1989, p. 365; et alii, 1996, p. 130), many species (mainly South American) are reported from the Late Albian (P. barbacoense, P. caneroti, P. discoides, P. duplicatum, P. elegans). Furthermore, in Europe and Asia it is also known from the Late Albian (P. zagrosiense). Based on our recent knowledge, the stratigraphical distribution of Parengonoceras spans the whole of the Albian.
Included species:
Remarks. In his accurate analysis, (1970, p. 1030-1032) examined the sutural development of Parengonoceras during ontogeny, proving that it has two distinct adventive lobes. Parengonoceras has the most complex suture lines among the Early and Middle Albian Engonoceratidae with irregularly and finely frilled deep lobes and saddles. This suture line is distinctive among other Early and Middle Albian engonoceratid ammonites and is considered to be the forerunner of the suture seen in Hypengonoceras and therefore, a phylogenetic connection exists between these genera. All species of Parengonoceras have rather compressed and smoothly ornamented shells (although fine tubercles may appear on flanks) with frilled suture, which clearly distinguish them from all other genera of Albian and Cenomanian engonoceratids. The proposed Parengonoceras - Hypengonoceras lineage and the suture remarkably different from those of other engonoceratid species suggest a polyphyletic origin for the Engonoceratidae.
Genus Platiknemiceras ,
1954, p. 174
= Platyknemiceras , 1959, p. 77
(Fig. 10 )
Type species. Platiknemiceras bassei , 1954, from the Albian of Spain.
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Figure 10: Platiknemiceras bassei , 1954. Reproduction of the figure of (1954, p. 175, refigured by , 1961).
Diagnosis. Diagnosis is after (1954, p. 175). Discoidal shape, flanks convex in the middle, being widest next to umbilicus, which is narrow. Umbilical wall is vertical, the last whorl almost entirely embracing. Sculpture is reduced consisting of attenuated ribs that arise at the umbilicus. Ribs may be flexuous in course but not tuberculate. Venter is smooth and forms a shallow groove. Suture is ceratitic.
Occurrence. Platiknemiceras appears in the Early Albian. Most species occur in Early-Middle Albian and some range into the Late Albian (P. deserti, P. jullieni) and higher, into the earliest Cenomanian (P. valencianum). Geographically the genus is reported from Europe (Spain, France), Africa (Algeria, Egypt), South America (Colombia, Peru), North America (Texas, USA), the Middle East (Iran) and Japan.
Included species:
Remarks. (1961) discussed the genus thoroughly and grouped some species of Knemiceras (K. deserti, K. gracile, K. hachourii, K. subcomplicatum, and K. sp. nov. [later described as P. flexuosum et alii, 1998a]), and Engonoceras (E. jullieni) under Platiknemiceras because of common morphological features: thinly discoidal shape with extreme reduction of ornamentation; this grouping restricts the scope of the genus Knemiceras. Platiknemiceras is close to Parengonoceras (, 1961, p. 354) but it includes only compressed and discoidal forms having extremely reduced sculpture. P. caseyi sp. nov. was listed previously (, 1954; & , 1964) as Engonoceras aff. stolleyi (), an attribution corrected by et alii (1980, p. 334). Since (1961) revised the genus there have been no further contributions regarding the content and systematics of the taxon; however four new species have been described. Most recently (2010, p. 171) discussed the species assigned of Platiknemiceras. He considers the genus to be restricted to P. bassei, P. hachourii, and P. flexuosum. (2010) transfers back to Knemiceras some of the species that (1961) referred to Platiknemiceras (P. gracile, P. subcomplicatum, and P. deserti). Unfortunately, (2010) did not consider the other five species (P. caseyi, P. colombiana, P. julieni, P. sequanense, and P. valencianum) of the genus so his revision is not complete. For the moment this genus is considered to be polyphyletic for it is based on morphological features common to all.
[Engonoceratidae gen. et sp. nov. ,
1973
= Borissiakoceras sp. ( & ,
2009, p. 881)]
This tiny specimen is reported from the Albian of New Zealand (, 1973, p. 106). The specimen is not strongly evolute (two whorls are visible) atypical for Engonoceratidae. Nor is the suture is typical of Engonoceratidae: therefore the generic assignment is not supported. & (1979, p. 116) had already referred it to Borissiakoceras, but without further analysis. This was provided by & (2009) who also assigned it to Borissiakoceras. However poor preservation and a limited number of specimens do not permit a more precise distinction.
The mean generic range of Jurassic/Cretaceous ammonites is 7.3 myr per ammonite genus ( & , 1983). Engonoceratid genera have longer ranges. The longevity of most of the genera is remarkable: seven genera existed for more than 10 myr (Engonoceras, Glottoceras, Knemiceras, Metengonoceras, Parengonoceras, Platiknemiceras, and Protengonoceras). Only two genera (Hypengonoceras and Neolobites) had shorter ranges (4 myr) than the average mean generic range. Originally the Engonoceratidae were said to have appeared during the Barremian (, 1938, p. 486), but later works did not support this view. Now it is widely accepted that engonoceratids appeared in the latest Aptian-earliest Albian (, 1956; , 1981; , 2010) in shallow marine platform environments of the southern Tethys (Algeria) and migrated to the peri-Tethyan territories (, 1957; , 1965). It is noteworthy that G. (1940b, p. 1066) had already speculated that some engonoceratids possibly were present in the Aptian of Texas, an observation supported by et alii (2004) when revising the original collection of Ferdinand , that included an Engonoceras sp. from the ?Upper Aptian of Texas. Obviously, the adaptive radiation of the group was rapid during the Early Albian, for its genera played an important role in the faunas of the Peruvian Basin (South America) and the Western Interior Sea (USA). During Middle and Late Albian times the group occupied the Far East and Australian provinces (Australia, Borneo, India, Japan, Madagascar and South Africa).
In the Cenomanian it successfully invaded some portions of the Boreal Realm (Mowry Sea, Canada) for it became ubiquitous in temperate watery environments everywhere in palaeolatitudes up to the 60 degrees where it co-occurs with Boreal ammonites (, 2009). In the Mowry Sea the adaptive radiation of engonoceratid ammonites was so successful that the invader Metengonoceras displaced the endemic Neogastroplites from its original morphoplace (, 2004) as recorded by changes in its morphotypes (, 2000). The taxon became extinct during the Cenomanian. The last surviving species of the engonoceratids are reported from the latest Cenomanian of Texas ( et alii, 1981), Niger and Nigeria ( et alii, 1992, 1994; et alii, 1998) and the southern Mediterranean ( & , 2005; , 2007; et alii, 2008). Table 1 summarizes the stratigraphical ranges of engonoceratid genera. Although (1935) reported a Metengonoceras dumbli from the Early Turonian of Niger, this dating is not supported by the detailed biostratigraphical study of et alii (1992), which showed that M. dumbli ranges only to the latest Cenomanian in Niger. The general climate warming of the mid-Cretaceous combined with an increase in subtropical upper ocean temperature ( et alii, 2003), may have helped the radiation of engonoceratids along ancient continental shelf seas. Changes in sea level may be an additional favourable factor. The highest relative sea level documented occurred during the Mesozoic ( et alii, 2003a). These fluctuations may have been accompanied by changes in the oceanic circulation that aided in increasing productivity and opened new ecological niches, which together resulted in a decline of provincialism in epicontinental seas. Some of these factors may have contributed to a general decline of ammonoid diversity ( et alii, 2003a, p. 395). However, the well documented decrease in the richness of ammonoid species significantly pre-dates the spread of anoxic deposits ( et alii, 2003b) and surely contributed to the dramatic changes of shallow water habitats that were preferred by engonoceratids, that in the end may have led to their extinction. "Suddenly" raised sea levels would have reduced the number of barriers between those epicontinental and shallow water habitats and endemic centres in which engonoceratids lived.
Based on our recent knowledge, all engonoceratid genera preferred shallow marine environments, and some (Knemiceras and Neolobites) occupied extremely shallow (tidal, lagoonal) milieux. However, it took a long time to arrive at this conclusion, which began seventy years ago.
G. (1940a) was among the first to discuss the growing importance of palaeoecology, pointing out that only oxyconic ammonites (as Engonoceras and Oxytropidoceras) are numerous in the epineritic area of the Cretaceous of Texas (Western Interior Sea, USA) where echinoids, oysters, and thick-shelled gastropods abundant. G. (1940a) estimated the epineritic zone to occupy water depths of 40 to 180 metres, which corresponds well with the lower limit of the euphotic zone. G. (1940b, p. 1066) considered that the group preferred shallow water and platform facies: "Knemiceras, Engonoceras and Oxytropidoceras appear to have had their greatest development in the broad, shallow, clear seas of Texas". The habitat-dependency of ammonites was also analyzed by (1991) for the Cenomanian-Turonian ammonites of the Western Interior. It became obvious that most ammonites are not restricted to a particular depth or facies, although compressed and discoidal forms (e.g. the Engonoceratidae) tended to occur in environments where the depth was less than 50 to 100 metres (, 2003). (1984) also analyzed thoroughly the Western Interior basin, its endemism and faunal provinces concluding that during the Cretaceous the maximum depth of thebasin was never greater than 200-500 metres with a significant area covered by less than a 50 metres. This shallow, warm, trophic environment offered excellent conditions for speciation among many ammonites (including engonoceratids) thus creating an endemic centre (, 1972). Based on his extensive research on bivalves, (1973) defined a Western Interior Endemic Centre (WIEC). Later et alii (1993) enlarged the scope of the WIEC to include other molluscs emphasizing that the WIEC was a very important pool of genetic variability and speciation for many molluscs. This direct link between the water depth and the blossoming of the engonoceratids is remarkable. Its validity supported by (1990, 1996) who from a theoretical standpoint pointed out the special ecological needs of engonoceratids and that they preferred shallow (30-50 metres) water. Ecology of the South American Engonoceratidae was first discussed by (1970, p. 1022) concluding that the South American Engonoceratidae lived on a subsiding carbonate platform. (1995a) also recognized the importance of engonoceratids in shallow marine facies: "Knemiceratinae (Knemiceras, Platiknemiceras, Parengonoceras) represent important ammonoid "pioneer" genera for mid-Cretaceous transgressive facies: they occur frequently near the base of marine, often glauconitic sediments overlying a continental clastic succession in their Tethyan (s.l.) realm." Engonoceratids frequently occur in glauconitic sediments. Parengonoceras duboisi and Knemiceras saadense are present in the glauconitic sand/sandstone of Salazac (Gard, France: , 1940; , 1989) and in the latest Albian glauconitic limestone of Pénzesgyor (Bakony Mts. Hungary; , 1990a, 1990b), furthermore Parengonoceras attenuatum spinosum of (1963) from the Early Albian of Bathurst Island (Australia) is found in a glauconite sand. et alii (1981) reported an engonoceratid (Metengonoceras dumbli) from the Late Cenomanian of stable Europe (Angers, NW France), also in a glauconitic sand. (1997) noted a Knemiceras from a glauconite bed in the Albian of Jordan. & (1981) described Parengonoceras caneroti from a lignitic deltaic series of latest Albian age at Castellon (Spain). This occurrence also supports the shallow water dependency of engonoceratids. Although & (1991) considered engonoceratid ammonites to be indicative of a neritic environment they did not discuss their shallow water dependency.
It became obvious very early on, that the presence of some engonoceratids may be indicative of certain environments. (1903, p. 148) called attention to the ostreid and bryozoid encrustations of Knemiceras syriacum shells. More recently (2008) too discussed shell encrustations on Knemiceras found in fluvial, deltaic, littoral or sublittoral facies. The encrustation may have occurred during post-mortem drifting, rather than during life in natural habitats. However, & (1976) and (1996) found that the post-mortem dispersal of ammonoids is rather the exception than the rule. This finding is very probably applicable to Knemiceras. Accepted as a generalization, nekroplanktonic dispersal on a large scale must be abandoned; a Knemiceras from the Albian of Jordan with oyster encrustation (, 2008) makes very likely the probability of a shallow water habitat. (1955) pointed out that the Knemoceras [sic!] beds are linked to basal conglomerate in Borneo. et alii (1986) remarked that various species of Knemiceras occur in a littoral facies of the southern Atlas Mountains, Africa. et alii (1997) report Knemiceras from a delta-influenced carbonate ramp in the Albian of Sinai (Egypt), again supporting the conclusion that many occurrences of Knemiceras are coincident with the early stages of transgressive systems tracts. & (2008) report Knemiceras from intercalated non-marine-shallow marine beds of Middle Albian age in Israel. Knemiceras uhligi from the Albian of Oman occurs in a shallow marine (average depth 35 metres) epicontinental sea, which might not have been more than 80 metres deep ( et alii, 2000, p. 223). But ( & , 1994; et alii, 2009) report that Late Albian 'Knemiceras beds' in Tunisia involve a unique range of ecological needs. (1956) began the attempts to explain the ecology of the highly variable Knemiceras. She postulated that as Knemiceras lived in a stable environment in the neritic zone at the margin of the old Saharan mainland where there was no deposition of detritus or perturbation, so conditions there were optimum for the rapid expansion of its population, and the stress that would have hindered adaptation did not exist. Consequently, the genetic potential and variability of Knemiceras was wasted in useless polymorphism. (1956) said that the mode of life of Knemiceras species was more benthic than neritic. She outlined a completely opposing view of the mode of life and habitats of Knemiceras and other engonoceratids. & (1991) proposed that the great variability of shell form and ornamentation can be explained by their inhabintance of a shallow and labile (= epicontinental) environment. Recently (2008) reported Knemiceras from the euphotic, shallow marine deposits of Late Albian age in the Zagros Mountain (SW Iran). Again it is the only ammonite found in that shallow marine environment. The shallow water dependency of Knemiceras is supported by the frequent presence of Knemiceras on shallow marine platforms ( & , 1994) and in fluvial-lagoonal deposits with dinosaur remnants ( et alii, 2000). Generally, Knemiceras is the only ammonite in shallow water environments of the Arabian Plate (, 2010). Dinosaur experts also remark that during Cenomanian times the fragmented dinosaur remains found in continental and deltaic facies followed by lagoonal and coastal plain deposits topped by marine limestones containing the first truly marine animals such as Neolobites ( et alii, 2005, p. 168). & (2005) noted that where Neolobites occurs in great abundance, it is on shallow-marine shelves. Neolobites is always linked to transgression and/or flooding periods in South-America ( & , 1993, p. 600). Distribution of Cenomanian carbonate platforms and Neolobites occurrences are correlated to a fine degree in northern Africa and the Americas ( & , 1993). The possible shallow marine dependency of Engonoceratidae is strongly supprted by (2007) who considers Neolobites as the endemic and intra-shelf taxon of the group. That Neolobites is a Cenomanian intra-shelf (e.g. shallow marine) taxon is strong support for the same mode of life for its predecessor, the Albian engonoceratids; however the Neolobites in Bolivia (, 1968; & , 1992) does not support its endemism at the genus level.
With regard to the dispersal of Ammonoidea, (1993) analyzed the post-hatching early life of 71 selected Cretaceous species of Ammonoidea. He concluded (based on density calculations of the shells), that at the hatching stage most of them were floaters with a positive buoyancy, strongly suggestive of a planktonic mode of life, thus permitting the rapid migration of all Cretaceous Ammonoidea. But positive buoyancy cannot explain the shallow water dependency of all engonoceratids, a relationship strengthened by 's (1988) report of well preserved and abundant engonoceratid fauna (Parengonoceras, Protengonoceras, Engonoceras pierdenalis, E. stolleyi, and E. uddeni) from the Middle Albian open shelf of Lampazos, Sonora (Mexico). (1981) offered an interesting working hypothesis to explain the connection between the distribution of engonoceratids and the depth of seas in which they occur. He noted that thin forms [of engonoceratids] are found in the western regions and the thickest in eastern areas. This difference would suggest some regional control in the depth of the seas. But recent knowledge of the geographical and facies distribution of engonoceratids, negates this idea. Thin forms are found in the Far East (Platiknemiceras caseyi in the Early Albian of Japan ( et alii, 1980).
Some Upper Albian ammonoid habitats in southeastern France were reviewed by et alii (2005). But this work did not discussed the habitats of the engonoceratids, although engonoceratids occur in that region (cf. , 1940; , 1989). Based on the available data engonoceratids certainly lived in the neritic and epipelagic domain of et alii (2005) occupying extremely shallow marine habitats. But we have more evidence for the connection between transgression and the rapid distribution of engonoceratids. The "pioneerism" of engonoceratids sensu (1995a) is linked to their shallow water habitancy, for the colonizing populations of engonoceratids were able to occupy rapidly the new areas of shallow water formed by transgressive seas, and thus to inhabit pristine ecological niches. The invasion of engonoceratids into the Peruvian province ( & , 2004) is closely linked to a transgression. In Early Albian times, the stem-engonoceratids which originated at the southern margin of the Tethys invaded the Peruvian Basin using shallow water seaways. A similar event occurred during the Cenomanian. (2007) following up on earlier works noted that the Cenomanian shallow water seaway between the southern margin of the Tethys and South America (Saharan seaway), Metengonoceras migrated southward during latest Cenomanian times (, 2007, fig. 1). (1935) and et alii (1992) had already reported this southward advance on the Late Cenomanian deposits of Niger. The appearance of M. oliveirai in Sergipe, Brazil (, 1967, 1986; , 1994) substantiates this southward migration of the seaway.
(1938, p. 510) attempted to erect a phylogeny of Engonoceratidae, proposing that the first engonoceratid was the Aptian Protengonoceras with no direct link to other engonoceratids. The first Albian engonoceratid genus was Engonoceras and its offsprings were Metengonoceras, Parengonoceras, Hoplitoides, Knemiceras, Neolobites, and Placenticeras. (1952) suggested that in all probability, the Engonoceratidae were derived directly from the Parahoplitidae. (1957, p. 39) proposed that if the pseudoceratitic development of the Albian species is indeed the expression of a development from the deshayesitid lineage, then the superfamily name Placenticeratoidea of (1900, p. 584) would be applicable. Summarizing the earlier results, (1960, p. 208) concluded: "origin of the Engonoceratidae has never been settled." Since that statement, there has been no significant development concerning the origin of the engonoceratids. Almost forty years later, the situation is unaltered from the state that et alii (1998a, p. 1) remarked: "evolutionary origin of the group is uncertain". Recently & (2001) outlined a phylogenetic chart for Engonoceratidae proposing that the most possible ancestor was Knemiceras ( & , 2001, fig. 2) with a questioned relation between Parengonoceras/Hypengonoceras and the rest of the group. So uncertainty remains an open field for investigation.
Speculation on the origin of Engonoceratidae and its possible parahoplitid origin now is strengthened by new results on the ecology of Parahoplites. et alii (2009, p. 904) report Parahoplites from a lagoonal environment in north-central Tunisia. In addition to the shallow marine habitat of Parahoplites, et alii (2005) proposed that the relationship of the mode of life of the Acanthohoplitinae to the water surface (incl. Parahoplites) supports the probability of a relationship. The similarities of the environments of Parahoplites and Engonoceratidae furnish a plausible explanation for the origin of the Engonoceratidae.
Recently, & (2004) and (2008) placed the Engonoceratidae in the superfamily Pulchelliatoidea based on morphological analogies. However, the ecological data on Pulchelliidae sensu stricto do not fully support this lineage. Recently et alii (2004) proposed a nektobenthic mode of life for the Pulchelliidae. If the Engonoceratidae is in fact derived from the Pulchelliidae, then their differences in modes of life and palaeoenvironments require an explanation. Other data ( et alii, 2008) also refer to neritic platform environment with an abundant pulchelliid fauna. Very recently (2010) rejected the pulchelliatoid origin for the Engonoceratidae.
A search for the evolutionary driving force behind the genesis of the Engonoceratidae suggests that the global oceanic anoxic event offers a plausible explanation. Dramatic changes in the pelagic ecosystem may have been triggered by the long anoxic event (OAE 1b) and progressively increased ecological pressure on those ammonite populations inhabiting shallow marine and platform environments. They were forced to occupy pristine extremely shallow ecological niches, previously unused. The evolution of the supposed parahoplitid ancestors continued apace and the first Engonoceratidae (Knemiceras, Platiknemiceras, Parengonoceras, and Protengonoceras) appeared in the latest Aptian-earliest Albian times in the shallow water environments of one or all three large shelves in North Africa, the proto-Gulf of Mexico, and the Caribbean Province.
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Figure 11: Stratigraphical distribution of engonoceratid genera and supposed phylogeny of Engonoceratidae. Data from (1938), (1960, 1961), & (1989), et alii (1996), & (2004). Cretaceous time-scale after et alii (2004). OAE 1b-d dating from et alii (2006), OAE 2 dating from et alii (1999). Seawater temperature curve from et alii (2003).
The origin of Engonoceratidae is almost certainly polyphyletic as indicated by the remarkable differences between the sutures of Parengonoceras
- Hypengonoceras and those of the remainder of the group, along with the apparently contemporaneous occurrences of these species in shallow marine environments in both Americas and North Africa. However, the enigma of the systematic position of Hypengonoceras, the most problematic genus of the Engonoceratidae may also be the result of a polyphyletic origin. &
(1983, p. 868) suggested that Hypengonoceras may be an off-shoot of an early Late Albian placenticeratid, but as seems more probable,
(1960, p. 208) proposed that Hypengonoceras was derived from Parengonoceras or some allied members of the Engonoceratidae. This uncertainty of origin is shown in
Fig. 11
by the discrete tracing of possible lineages. The most probable course of evolution was from ?Parahoplites to Protengonoceras. Evolution from Protengonoceras toward Engonoceras (or from Engonoceras toward Protengonoceras proposed by & ,
2001) and Metengonoceras and ultimately Neolobites is reasonable based on shell morphology and the simplification of suture over time. Lately,
(2010, p. 169) supports this concept assembles these genera into a phylogenetic lineage. The other lineage is more speculative; however &
(2004) suggested that the South American Engonoceratidae had a platiknemiceratid origin. But this is not a solution applicable to the other species, for a direct lineage from Parengonoceras to Hypengonoceras is feasible. Many have speculated on the existence of a direct phylogenetic connection between the Engonoceratidae and the Placenticeratidae (,
1930, p. 390; et alii,
1957, p. L109), but
(1960, p. 208) and &
(1989, p. 365) negated the possibility that the Engonoceratidae was the direct ancestor of the Placenticeratidae. Therefore the proposed phylogenetic connection with Hypengonoceras as an off-shoot of an early placenticeratid ( & ,
1983) must be abandoned (Fig. 11 ). Today's opinion: the Engonoceratidae became extinct without descendants in latest Cenomanian times in the Western Interior Province (USA). However the dating of the end of the Cenomanian is debated by
R.W. et alii
(2009), as is the closing of the Saharan seaway (Niger and Nigeria).
The apparent similarities between the engonoceratid and ceratitid suture lines tempted palaeontologists to speculate on the cause of the repetition of similar structures (cf. , 1914, p. 30 on the Neolobites suture line). Recently et alii (2005) analyzed the ecology and immigration patterns of approximately 500 specimens of ceratitid Ammonoidea found in strata of the Late Anisian to Early Ladinian age in Germany. Their study was based on the morphological variability of ammonoid shells. The Germanic Basin was the site of a shallow sea during most of the Anisian and Ladinian ( et alii, 2005, p. 19). It was occupied repeatedly by incursions of ammonoids, crinoids and brachiopods coincident with intervals of high sea level. After each of these immigrations new species of Ceratites evolved ( et alii, 2005, p. 25) just as we had seen among Cretaceous engonoceratids in the Peruvian Basin and in the Western Interior Sea (USA). Cretaceous species of the engonoceratids exhibit remarkable morphological convergence very likely that of the Triassic species used in the ecological analysis of et alii (2005). The repetition of suture lines, external shell morphologies, together with records of a shallow marine habitat, successive migrations, and successful adaptation in both Middle Triassic and mid-Cretaceous forms suggest that this pattern is a very successful ammonoid eco-morphotype in shallow marine habitats. This conclusion is supported by the morphological convergences of the Triassic Ceratites nodosus nodosus ( et alii, 2005, fig. 4) and the Cretaceous Knemiceras syriacum (, 1848, figs. 2-3).
Other morphological convergences occur (Fig. 12 ) among the Triassic Discoceratites semipartitus ( et alii,
2005, fig. 4), the Albian-Cenomanian Metengonoceras teigenense ( & ,
1989, pl. 2, figs. 1-2) and the Maastrichtian Sphenodiscus pleurisepta ( et alii,
2005, fig. 7a-b). The Triassic scheme of colonization followed by speciation follows the same pattern as that of engonoceratids during the Albian. The endemic centres in the Americas were rapidly colonized by engonoceratids, followed by speciation ( & ,
2004). This suggests that the repetition of the Triassic eco-morphotype in the engonoceratids was caused by their occupation of nearly identical ecological niches and habitats.
Another interesting convergence is that of the engonoceratids and another group of Ammonoidea, the Maastrichtian sphenodiscids.
et alii (2005) reported several species of Sphenodiscus (S. lobatus and S. pleurisepta) from the Maastrichtian of La Popa Basin, Mexico, a shallow water coastal deposit. The oxyconic Sphenodiscus was well adapted to continuous swimming with excellent acceleration skills ( & ,
1996, p. 211). Shell morphology, suture lines and palaeoenvironments of these sphenodiscid ammonites surprisingly repeat that of the Albian-Cenomanian engonoceratids. et alii
(2005, p. 62) concluded: "sphenodiscid-dominated faunas […] are thus clearly related to bathymetry, i.e. the shallow-water assemblages are very low in diversity and represent specialization to an environment in which ammonoids could not easily survive". It seems obvious that during ammonoid evolution, the combination of shallow water habitats, the oxy- and/or platyconic, brevidome shell morphology and simplified, Ceratites-like sutures occurred at least three times, one of them the Albian-Cenomanian engonoceratids. In all cases, this pattern was associated with successful migration, colonization and speciation. This set of characters is summarized in
Fig. 12 .
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Figure 12: Strong similarities in shell shapes among certain ammonoid taxa. Repeated eco-morphotype pattern among Mesozoic Ammonoidea. The proposed eco-morphotype comprises a set of features consists of nektonic, active swimmer mode of life in shallow marine settings, simplified and ceratitic suture and involute, oxycone-platycone shell form. Mesozoic time-scale after et alii, 2004.
Engonoceratid ammonites are important time indicators of chronostratigraphic time in strata of shallow marine origin ranging in age from the ?latest Aptian times to the youngest of Cenomanian formations. Where they occur in the Tethyan Faunal Realm, they are excellent indicators of a shallow water environment. Today engonoceratids play an important role in correlation, biostratigraphy and ecological studies, but trhey are limited in distribution to temperate water. Engonoceratids are a versatile, and successful group of Cretaceous Ammonoidea, ecological pioneers that were successful in occupying previously inaccessible pristine ecological niches. The range of suture lines, shell form, all is response to a shallow water habitat created a morphological pattern that proved successful not only for the engonoceratids, but also for other Mesozoic Ammonoidea.
Engoncoeras duboisi , 1989, is known only from 2 localities and represented by a handful specimen. New specimen from Hungary led the present author for thorough morphological analysis, which resulted the below revision of the species.
Parengonoceras duboisi (, 1989)
(Figs. 13A-B
- 14
- 15 )
v 1989 Engonoceras duboisi sp. nov. : p. 56, pl. 2, figs. 1-3.
1989 Engonoceras sp. cf. E. duboisi sp. nov. : p. 57, pl. 2, fig. 4.
v 1990a Engonoceras duboisi ; : p. 11, pl. 1, figs. 1-4.
v 2007 Engonoceras duboisi ; et alii: p. 101, pl. 15, figs. 1, 4.
A |
 |
B |
 |
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Figure 13: Parengonoceras duboisi (, 1989) from the latest Albian (Stoliczkaia dispar Zone) of Tilos forest (Bakony Mts. Pénzesgyor, Veszprém County, Hungary). Specimen NHNM M2010.760.1. A.) lateral view, B.) dorsal view.
Click on thumbnail to enlarge the image.
Figure 14: Suture line of Parengonoceras duboisi (, 1989) at whorl height of 51 mm. Specimen HNHM M2010.760.1.
Material. Three internal moulds from the Late Albian (Stoliczkaia dispar Zone) of Tilos forest (Pénzesgyor, Bakony Mts, Hungary) and three worn internal moulds from the Late Albian (Stoliczkaia dispar Zone) of Salazac, Gard (France).
Occurrence. (1989) reported it from the latest Albian of Salazac, Gard (France). (1990a) described it from Hungary (Tilos Forest, Bakony Mts.). et alii (2007) copied the figures of (1990a). New collection by the present author from the Tilos Forest (Bakony Mts., Hungary) yielded a single specimen described here for the first time supporting the distribution of the species for the latest Albian of France and Hungary. The new record of the Late Albian P. duboisi from the Bakony Mts. is important as a connector between the rich and diverse western and less well known eastern Tethyan Parengonoceras species during the Albian. It suggests a southern Tethyan migration route because the fact that the Alcapa Tectonic Unit (which also includes the Bakony Mts.) was part of the Apulia microplate bordered by the southern edge of the Penninic Ocean ( & , 2004). The revealed third specimen of P. duboisi from Hungary refers to the rarity of this species in the Bakony Mts, Hungary: 1:10,000 (Z. Evanics pers. comm.).
Description. Medium sized incomplete but shelly phragmocones and internal moulds. Coiling is typical of the Engonoceratidae. The conch is discoidal and strongly involute and oxycone
(Fig. 13 ). Last whorl embraces the previous ones. The umbilicus is small and shallow. Umbilical wall is low and the umbilical shoulder is gently rounded. Flanks converge and are almost unsculptured. The cross section is lanceolate. Venter is smooth and narrow. Finer or stronger prorsiradiate falcoid ribs arise at midflank, which end at the ventro-lateral shoulders. Six fine and falcoid ribs are on the last quarter of whorl; ribs end at fine tubercles on ventral shoulder. Body chamber is not preserved. Suture line
(Fig. 14
) is typical for Parengonoceras showing two adventive lobes. Suture is characterized by pincer-like and finely incised lobes and saddles. Pincer-like saddles are irregular polyphillic. Lobes are also incised irregularly.
Click on thumbnail to enlarge the image.
Figure 15: Size distribution of Hungarian (solid) and French (open) specimens of Parengonoceras duboisi (, 1989).
Remarks. P. duboisi is most similar to P. elegans and Platiknemiceras hachourii (). The Early Albian Platiknemiceras hachourii has a slightly more slender cross-section, and the striae on the lower flanks are finer in P. duboisi and the suture is less frilled than on P. hachourii. The extreme reduction of ornamentation in P. duboisi is also not present in P. hachourii. The ornamentation is typical of Parengonoceras as is the diameter and whorl height ratios
(Fig. 15 ). P. duboisi is most similar to P. elegans in its shell ornamentation. The small, ventrolateral falcoid ribs of
P. elegans (,
1970, pl. 3, fig. 1a-c) are comparable with P. duboisi. However P. duboisi shows fewer ribs per whorl than P. elegans. The superficially similar Engonoceras and Parengonoceras species are clearly distinguished from each other based on suture lines
- if present. Normally the saddles of Engonoceras are all entire except that the outermost, which is bifid ( et alii,
1996, p. 130). The present species show only frilled saddles in all stages in the specimens. All saddles of the holotype are incised, mostly bifid (,
1989, pl. 2, figs. 2-3) being worn specimens, therefore the frilled suture is not well preserved, however there are no entire saddles at any stage. The Hungarian specimens are well preserved, partly shelly specimens with well observable suture revealing more incised saddles. Biometry of the French and Hungarian specimens refers the same species
(Fig. 15 ), too. Based on the well developed and frilled saddles and lacking entire saddles at any stage, this species is convincingly placed into Parengonoceras. Recently the uncertainty of the systematic position of P. duboisi is also referred by
(2010, p. 169) excluding it from the genus Engonoceras. This conclusion is supported here.
I acknowledge the valuable comments on the initial draft by Herbert , Jean-Louis and Christian . My special thanks to Robert W. for providing vital papers and essential criticism of draft as well as to Peter F. to clarify 'Pseudengonoceras'. I acknowledge the detailed remarks of Jens , which greatly improved the text. Special thanks to Margaret , Jeffrey and Mikel for sending important papers. I am deeply indebted for the great help and encouragement of Hugh G. on Albian ammonites found in the 1990's and especially for his careful comments on this draft, which greatly improved the quality of the paper. I am also indebted to the late William who in the early 1990's introduced me to the engonoceratids. I acknowledge the technical support of Attila (Hungarian Natural History Museum, Budapest) for housing the specimen treated here and granting permission for use the facilities of the Museum. I am grateful to Mr. Zoltán (private collector) who provided his Hungarian engonoceratid specimens for this research. Finally, I acknowledge the comments of the referees of this paper, especially the detailed remarks of Christian . Last but not least I acknowledge the continuous help of the Editor, Bruno , and the careful linguistic improvement provided by Nestor .
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| Species | Stratigraphical distribution | Geographical distribution | References | |
| Glottoceras , 1875 | Albian | South America | ||
| 1 | andinum (, 1970) | Upper Albian | Venezuela | , 1970 |
| 2 | attenuatum , 1875 | ?Cenomanian | Celendín, Peru | , 1903; , 1938 |
| Albian-Cenomanian | Oriente Basin, Ecuador | & , 1998 | ||
| Albian | Une, Une Formation, Colombia | , 2002 | ||
| Middle Albian | northern Peru | et alii, 1947; , 1956 | ||
| Lower Albian | Peru | et alii, 2003; & , 2004 | ||
| 3 | bassleri ( in et alii, 1947) | Aptian | En Huanco, Celendin, Peru | et alii, 1947 |
| 4 | crassicostatum (, 1910) | Middle Albian | Peru | et alii, 1947 |
| Lower-Middle Albian | Pongo de Rentemo, Peru | et alii, 1998 | ||
| 5 | crassinodosum (, 1910) | Middle Albian | Condebamba, northern Peru | , 1910; et alii, 1947 |
| Lower-Middle Albian | Peru | et alii, 2003; & , 2004 | ||
| 6 | gabbi (, 1903) | ?Cenomanian | Quebrada de Huari, Peru | , 1903; , 1938 |
| Middle Albian | Celendín, Sunchubamba, northern Peru | , 1956 | ||
| upper Lower Albian | Peru | et alii, 2003; & , 2004 | ||
| 7 | gracilecostatum (, 1910) | Lower Albian | northern Peru | et alii, 2003 |
| 8 | laraense (, 1970) | Middle Albian-Upper Albian | Misahualli, Ecuador; Venezuela | et alii, 2007 |
| 9 | largum , 2002 | Lower Albian | La Cuenca, Peru | et alii, 2003 |
| 10 | libertadense (, 1952) | Albian | Colombia | , 1952 |
| 11 | moorei ( in et alii, 1947) | upper Middle Albian | Peru | et alii, 2003; & , 2004 |
| Middle Albian | Peru | et alii, 1947 | ||
| Misahualli, Ecuador | et alii, 2007 | |||
| 12 | ollonense (, 1877) | Lower Albian | La Cuenca, Peru | et alii, 2003 |
| Sunchubamba, northern Peru | , 1956; et alii, 1998 | |||
| 13 | ovale (, 1956) | Middle Albian | Uchupata, northern Peru | , 1956 |
| 14 | pacificum (, 1956) | Lower Albian | Celendín, Peru | , 1956 |
| 15 | pegnai (, 1979) | Cretaceous | Colombia | , 1979 |
| 16 | raimondii (, 1908) | Middle Albian | Cajamarca, Sihuas, Pomachaca, northern Peru | , 1910; et alii, 1947; , 1956; et alii, 1998 |
| upper Lower Albian | Peru | et alii, 2003; & , 2004 | ||
| 17 | semicostatum (, 1910) | Middle Albian | Cochamarca, northern Peru | , 1910; et alii, 1947 |
| upper Lower Albian | Peru | et alii, 2003; & , 2004 | ||
| 18 | seminodosum (, 1910) | Middle Albian | Peru | & , 2004 |
| 19 | sommermeieri ( in et alii, 1947) | Middle Albian | Peru | et alii, 1947 |
| Misahualli, Ecuador | et alii, 2007 | |||
| 20 | spinosum (, 1910) | Middle Albian | northern Peru | et alii, 1947 |
| Lower Albian (mammillatum Zone) | Bathurst Island, North Australia | , 1963 | ||
| 21 | tardum (, 1956) | Middle Albian | Sihuas, northern Peru | , 1956 |
| Lower Albian | La Cuenca, Peru | et alii, 2003 | ||
| 22 | triangulare (, 1956) | Middle Albian | Sihuas, northern Peru | , 1956 |
| 23 | typicum (, 1910) | Middle Albian | northern Peru | & , 2004 |
| Engonoceras & , 1881 | Lower Albian-Middle Cenomanian | Tethyan | ||
| 1 | belviderense (, 1894) | Lower Cretaceous | Belvidere, Kansas | , 1903; , 1987 |
| Upper Albian | Texas, USA | et alii, 2003 | ||
| Neocomian | Kansas, USA | , 1970; et alii, 1998a | ||
| Middle-Upper Albian | Sonora, Mexico | , 1938; & , 1991 | ||
| 2 | commune , 1903 | Cretaceous | Texas | , 1903 |
| 3 | complicatum , 1903 | Albian | Texas | , 1903; et alii, 1998a |
| Upper Albian | El Mizab, Meknes, Atlas, Morocco | & , 1968 | ||
| 4 | elegans et alii, 1998a | Albian | Texas | et alii, 1998a |
| 5 | emarginatum (, 1893) | Albian | Pleasant Point, Texas, USA | , 1903 |
| 6 | gibbosum , 1903 | Albian | Texas, USA | , 1903; et alii, 1998a |
| Middle Albian | et alii, 2003 | |||
| 7 | grimsdalei , 1931 | Upper Albian | Weymouth, Dorset, England | , 1931; , 1956 |
| 8 | hilli , 1898 | Albian | Texas, USA | et alii, 1998a |
| North Peruvian Andes | et alii, 2003 | |||
| Middle-Upper Albian | Texas, USA | et alii, 2003 | ||
| 9 | iris , 1924 | Middle Albian | Folkestone, England | , 1938; , 1924 |
| 10 | jezzinense , 1940 | Albian | Djebel Tebega, Tunisia | , 1956 |
| Jezzine, Lebanon | , 1940 | |||
| 11 | khenchelaensis (, 1940) | Albian-Cenomanian | Syria | et alii, 1998a |
| Albian | Bhamdoum, Lebanon | , 1961 | ||
| 12 | olgaluciae , 1979 | Middle Albian | Colombia | , 1979 |
| 13 | pierdenalis (, 1848) | Upper Cretaceous | Fredericksburg, Texas | , 1848 |
| Albian | Texas | , 1903; , 1938 | ||
| Upper Albian | Chihuahua, Mexico | , 1910 | ||
| Middle Albian | Sonora, Mexico | , 1988; & , 1991 | ||
| 14 | pinax (, 1902) | Cenomanian | Temojoh, West Borneo | , 1902; , 1940; , 1955 |
| 15 | retardum , 1903 | lowest Cenomanian | Pottsboro, Texas, USA | , 1903; & , 1989; et alii, 2005 |
| 16 | serpentinum (, 1900) | Lower Cenomanian | Texas | , 1982 |
| Upper Albian | , 1938; et alii, 1996 | |||
| uppermost Albian (dispar Zone) | , 1903; et alii, 1998b; , 2004; et alii, 2005 | |||
| 17 | subjectum , 1903 | uppermost Albian (dispar Zone) | Texas | , 1903; et alii, 1998b |
| 18 | stolleyi , 1898 | Albian | Texas | et alii, 1998a |
| Upper Albian | Chihuahua, Mexico | , 1910 | ||
| Middle-Upper Albian | Sonora, Mexico | & , 1991 | ||
| Middle Albian | , 1988 | |||
| La Cuenca, Peru | et alii, 2003 | |||
| 19 | thomasi , 1907 | Cenomanian | Rekkame, Middle Atlas, Morocco | & , 2009 |
| U. Albian-L. Cenomanian | Guelta, Tunisia | , 1938; , 1989 | ||
| 20 | toussainti , 1907 | U. Albian-L. Cenomanian | Tebaga, Tunisia | , 1938; , 1989 |
| 21 | uddeni (, 1900) | Comanche series, Kiowa shale | Kansas, USA | , 1903 |
| Middle Albian | Sonora, Mexico | , 1988 | ||
| 22 | vicorpense (, 1940) | Albian-Cenomanian | Syria | et alii, 1998a |
| Upper Albian | Djebel Naimia, Chotts, Tunisia | et alii, 2008b | ||
| 23 | wilkinsoni , 1956 | ?Upper Albian | Oregon, USA | , 1956 |
| 24 | zumoffeni (, 1940) | Albian-Cenomanian | Syria | et alii, 1998a |
| Neolobites , 1882 | Cenomanian | Tethyan | ||
| 1 | bassleri , 1926 | Cenomanian | Caminaca, Peru | , 1926; & , 1936 |
| Middle-Upper Cenomanian | Altiplano, Peru | , 1995; et alii, 1998 | ||
| 2 | brancai & , 1908 | U. Cenomanian- L. Turonian | Egypt | & , 1908; , 1914; , 1928 |
| Upper Cenomanian | Central High Atlas, Morocco | & , 2002 | ||
| Upper Cenomanian | Gafsa, Central Tunisia | & , 2005 | ||
| 3 | bussoni , 1965 | Cenomanian | Tinrhert, Sahara, Algeria | , 1965; et alii, 1981 |
| Upper Cenomanian | Ourém, Portugal | & , 1978, 1979 | ||
| 4 | choffati , 1903 | Cenomanian | Portugal | , 1903 |
| 5 | fourtaui , 1907 | Cenomanian | Egypt, Tunisia | , 1907; , 1915 |
| Upper Cenomanian | Tarfa, Arabic Desert | , 1928 | ||
| Tinrhert, Sahara, Algeria | , 1965; et alii, 1981 | |||
| Middle-Upper Cenomanian | Palmyrids, Central Syria | et alii, 2003 | ||
| 6 | isidis , 1915 | Cretaceous | Egypt | , 1915 |
| Upper Cenomanian | Gafsa, Central Tunisia | & , 1996 | ||
| 7 | kummeli , 1956 | base Upper Cenomanian | Celendín, Northern Peru | , 1956 |
| Bolivia | , 1968; & , 1992 | |||
| 8 | maresi (, 1862) | Cenomanian | Egypt | , 1862; , 1915; & , 1984 |
| 9 | medeninensis , 1954 | Upper Cenomanian | Tunisia | , 1954; & , 2005 |
| 10 | peroni , 1903 | Upper Cenomanian | Tinrhert, Sahara, Algeria | , 1910; , 1965; et alii, 1981 |
| Cenomanian | North Africa | , 1903 | ||
| Egypt | , 1914; , 1928 | |||
| Provence (France), Tunisia | , 1938 | |||
| 11 | pervinquieri & , 1908 | Upper Cenomanian | Egypt | & , 1908; & , 2005 |
| schweinfurthi & , 1908 | U. Cenomanian-L. Turonian | Egypt | & , 1908; , 1914; , 1928 | |
| Turonian | Egypt | , 1910 | ||
| 12 | vibrayeanus (d', 1841) | Cretaceous | Vibraye, Sarthe, France | d', 1841; , 1848 |
| Cenomanian | Celendín, Northern Peru | , 1926 | ||
| El Kisba, High Atlas, Morocco | & , 2009 | |||
| France | , 1942; et alii, 1996 | |||
| France, Portugal | , 1903 | |||
| Upper Cenomanian | Jordan | et alii, 2008 | ||
| Central Oman | & , 1991 | |||
| Estremadure, Lisbon, Portugal | et alii, 1979 | |||
| Vouillac, France | et alii, 1983 | |||
| Tinrhert, Sahara, Algeria | , 1965; et alii, 1981; et alii, 1996 | |||
| Egypt, Jordan | & , 2005 | |||
| Gafsa, Central Tunisia | & , 2005 | |||
| Tanout, Niger | et alii, 1992 | |||
| Burgos, Spain | , 1979 | |||
| Knemiceras , 1898 | ?Upper Aptian-Albian | Tethyan | ||
| as Knemiceras sp. | Upper Albian | Río Aguamulac, Huescar, Spain | et alii, 1982 | |
| Middle Albian | Aube, France | & , 1984 | ||
| 1 | aegyptiacum , 1955 | Upper Albian-Lower Cenomanian | Kazhdumi, southern Iran | , 1981 |
| Upper Albian | Gafsa, Central Tunisia | et alii, 2008 | ||
| Oued Zitoun, North Tunisia | & , 1986 | |||
| 2 | arambourgi , 1940 | Cretaceous | Syria, Lebanon | , 1940; & , 1991 |
| 3 | collignoni , 1955 | Cretaceous | Sinai, Egypt | , 1955 |
| Upper Albian-Lower Cenomanian | Kazhdumi, southern Iran | , 1981 | ||
| 4 | compressum , 1903 | ?Cenomanian | Beirut, Lebanon | , 1903; , 1938 |
| Albian | Nar, Kartang, Jahrum, southern Iran | , 1981 | ||
| Upper Albian | Venezuela | , 1970 | ||
| Oued Zitoun, North Tunisia | & , 1986 | |||
| Djebel Naimia, Chotts, Tunisia | et alii, 2008b; & , 2008 | |||
| Middle Albian | Bilbao, northern Spain | et alii, 2009 | ||
| 5 | douvillei , 1940 | Cretaceous | Djebel Dourra Syria | , 1940 |
| Upper Albian-Lower Cenomanian | Kazhdumi, southern Iran | , 1981 | ||
| Upper Albian | Lurestan, Iran | , 2007 | ||
| 6 | dubertreti , 1940 | Cretaceous | Djebel Dourra, Syria | , 1940 |
| Albian | Mderedj, Lebanon | , 1961 | ||
| Upper Albian | Lurestan, Iran | , 2007 | ||
| Lower-Middle Albian | Central Oman | & , 1991 | ||
| 7 | flexiloculosum , 1940 | Cretaceous | Syria | , 1940 |
| Middle Albian | Dead Sea, Israel | , 2002 | ||
| Lower Albian | Guajira, NE Colombia | , 2002 | ||
| 8 | iraniense , 1981 | U. Albian-L. Cenomanian | Kazhdumi, southern Iran | , 1981 |
| Middle-Upper Albian | Bangestan, SW Iran | , 2010 | ||
| 9 | kazhdumiense , 1981 | U. Albian-L. Cenomanian | Kazhdumi, southern Iran | , 1981 |
| Middle-Upper Albian | Bangestan, SW Iran | , 2010 | ||
| 10 | orientalis , 1955 | Middle Albian | Moghara, Egypt | , 1955 |
| 11 | persicum , 1981 | U. Albian-L. Cenomanian | Kazhdumi, southern Iran | , 1981 |
| Lower Albian | Bangestan, Lurestan, Iran | , 2007, 2010 | ||
| 12 | rittmanni , 1955 | Middle Albian | Moghara, Egypt | , 1955; & , 1955; , 2001 |
| 13 | saadense ( & , 1890) | Cretaceous | Tunisia | & , 1890 |
| Upper Albian | Salazac, Gard, France | , 1940; , 1989; , 2008 | ||
| Algeria | , 1910; , 1938; , 1941 | |||
| 14 | saharae , 1965 | Albian | Tebaga de Kebili, South Tunisia | , 1965 |
| Sinai, Egypt | et alii, 1997 | |||
| 15 | ? sinaiticum | Upper Albian | Gafsa, Central Tunisia | et alii, 2008 |
| 16 | spathi , 1955 | U. Albian-L. Cenomanian | Nar, Gavbasht, southern Iran | , 1981 |
| Upper Albian | Oued Zitoun, North Tunisia | & , 1986 | ||
| Lower Albian | northern Sinai, Egypt | et alii, 1997; , 2001; , 2008 | ||
| Bangestan, SW Iran | , 2010 | |||
| 17 | subcompressum , 1903 | ? Cenomanian | Gilead Mts. Jordan; Mt. Lebanon, Syria | , 1903; et alii, 1997 |
| 18 | syriacum (, 1848) | Lower Cretaceous | Lebanon | , 1848 |
| Angola | , 1931; , 1938 | |||
| Cenomanian | Mt. Lebanon, Syria | , 1903 | ||
| U. Albian-L. Cenomanian | Kazhdumi, southern Iran | , 1981 | ||
| Upper Albian | Portugal | , 1938 | ||
| Lurestan, Iran | , 2007 | |||
| Syria | et alii, 1996; et alii, 2003 | |||
| Middle-Upper Albian | northern Sinai, Egypt | et alii, 1997 | ||
| Jawan area, NW Iraq | , 2006 | |||
| Middle Albian | Tebaga, central Tunisia | et alii, 1985 | ||
| Moghara, Egypt | , 1916, 1928; & , 1955 | |||
| Dead Sea, Israel | , 2002 | |||
| Cajamarca, Uchupata, Peru | , 1956; et alii, 1998 | |||
| Lower Albian | northern Sinai, Egypt | , 2008 | ||
| 19 | uhligi (, 1886) | ? Cenomanian | Portugal | , 1903; , 1938 |
| Upper Albian | Santander, Cantabria, Spain | , 1996 | ||
| Teruel, Valencia, Spain | , 1995a, 1995b | |||
| Estremadure, Lisbon, Portugal | , 1941; et alii, 1979, , 1981 | |||
| Salinas (Angola); Moghara (Egypt) | , 1928, 1931 | |||
| Lurestan, Iran | , 2007 | |||
| Middle-Upper Albian | northern Sinai, Egypt | et alii, 1997 | ||
| Tebaga, central Tunisia | , 1965 | |||
| Muscat, Oman | et alii, 2000 | |||
| Lower Albian | Lurestan, Iran | , 2007 | ||
| northern Sinai, Egypt | , 2008 | |||
| Protengonoceras , 1903 | Lower/Middle Albian-?Lower Cenomanian | Tethyan | ||
| 1 | dufaurei , 1981 | Lower Cenomanian | Kazhdumi, southern Iran | , 1981 |
| 2 | gabbi (, 1898) | Albian | Sonora (Mexico), Texas | , 1903; et alii, 1998a |
| Upper Albian | northern Sinai, Egypt | , 2008 | ||
| Middle Albian | Texas | et alii, 1996 | ||
| Peru | et alii, 1998 | |||
| 3 | planum , 1903 | Upper Cretaceous | Dallas County, Texas | , 1903; , 1942 |
| 4 | ? prestati , 1981 | U. Albian-L. Cenomanian | Kazhdumi, southern Iran | , 1981 |
| Metengonoceras , 1903 | Middle Albian-Cenomanian | cosmopolitan | ||
| as Epengonoceras sp. | Upper Albian | Lower Saxony Basin, Germany | , 2007 | |
| as Metengonoceras sp. | uppermost Cenomanian | Sahara, Africa | , 2007 | |
| lower Upper Cenomanian | Sayreville, NJ, USA | & , 1990 | ||
| 1 | acutum , 1903 | Upper Cenomanian | North Central Texas | , 1903; , 1942; , 1988; & , 1990; et alii, 2003 |
| 2 | ambiguum , 1903 | Lower Cretaceous | Texas, USA | , 1903 |
| Albian | et alii, 1998a | |||
| Aptian-Albian | , 1938 | |||
| 3 | arnaudi , 1912 | Cenomanian | Touvois, Vendée, France | , 1942 |
| Charentes, France | et alii, 1981 | |||
| 4 | aspenanum ( & , 1931) | Cenomanian | Montana, Wyoming, USA | & , 1989; , 2004 |
| uppermost Albian | Montana, USA | et alii, 2003 | ||
| 5 | bedoti (, 1912) | Lower Cenomanian | Vendée, France | , 1942; et alii, 1981; & , 1984; et alii, 1992 |
| 6 | bravoense (, 1928) | Cenomanian [M. (P.) rhacioformis Zone] | Texas | , 1979 |
| lowermost Cenomanian | Texas, USA | et alii, 2005 | ||
| Albian-Cenomanian | Oriente Basin, Ecuador | & , 1998 | ||
| 7 | ? dibbleyi | Upper Cenomanian | Ouled nail, Saharan Atlas, Algeria | , 1999; et alii, 2008 |
| 8 | douvillei , 1912 | Cenomanian | Vendée, France | et alii, 1981 |
| 9 | dumbli (, 1893) | Cenomanian | Texas, USA | , 1942; et alii, 1996 |
| uppermost Cenomanian | Jordan | et alii, 2008 | ||
| Benue trough, Nigeria | et alii, 1998 | |||
| Upper Cenomanian | Mts. Iguellala, Niger | et alii, 1992 | ||
| Sinai, Egypt | & , 2001 | |||
| Le Mans, France | & , 1984 | |||
| Angers, France | et alii, 1981 | |||
| Middle-Upper Cenomanian | Gafsa, Central Tunisia | & , 2005 | ||
| Colorado, USA | & , 1972 | |||
| North Central Texas | & , 1990; et alii, 2003 | |||
| 10 | inscriptum , 1903 | Lower Cretaceous | Texas, USA | , 1903 |
| Upper Albian | et alii, 1996 | |||
| Middle Albian | & , 1989 | |||
| Albian | et alii, 1998a | |||
| Aptian-Albian | , 1938 | |||
| 11 | nigeriensis , 1935 | Upper Cenomanian | Niger | et alii, 1981 |
| 12 | oliveirai , 1967 | Albian-Cenomanian | Rio Fundo, Sergipe, Brazil | , 1986; , 1994 |
| 13 | teigenense & , 1989 | Cenomanian | Montana, USA | & , 1989; , 2004 |
| Lower Cenomanian | Normandie, France | et alii, 2002 | ||
| U. Albian-lowermost Cenomanian | Montana, USA | et alii, 2003 | ||
| 14 | tolveiense , 1912 | Cenomanian | Vendée, France | et alii, 1981 |
| Hypengonoceras , 1922 | Upper Albian | Tethyan | ||
| as Hypengonoceras sp. | Upper Albian | Makhtesh Ramon, Israel | , 1981 | |
| as Hypengonoceras sp. ind. | Upper Albian | Salazac, Gard (France) | , 1989 | |
| 1 | chouberti , 1966 | Upper Albian | Tarfaya, Morocco | , 1966; & , 1989 |
| 2 | decaryi , 1963 | Upper Albian | Madagascar | , 1963 |
| Zululand, Mozambique | , 1975; & , 1979 | |||
| Zufia, Navarra, Spain | , 1979 | |||
| 3 | fauremuretae , 1966 | Upper Albian | Tarfaya, Morocco | , 1966; & , 1989 |
| 4 | ? ibericum & , 1977 | Albian | Albacete, SE Spain | & , 1977; & , 1989 |
| 5 | tarfayense , 1966 | Upper Albian | Tarfaya, Morocco | , 1966; & , 1989 |
| 6 | warthi (, 1895) | Albian | Albacete, SE Spain | & , 1977 |
| Upper Albian | South India | , 1960; et alii, 1996 | ||
| Diego Suarez, Madagascar | , 1960 | |||
| Parengonoceras , 1924 | Albian | Tethyan | ||
| as Parengonoceras sp. juv. | Upper Albian | Suriname | & , 2006 | |
| as Parengonoceras sp. | Lower Albian | SW Iran | , 2010 | |
| as Parengonoceras sp. | Lower Albian | Kent, England | , 1957, p. 42 | |
| 1 | algerianum (, 1953) | lowermost Albian | Mellégue, Algeria | & , 2004 |
| 2 | azlense (, 1940b) | Upper Albian | Fort Worth, Texas | , 1940b |
| 3 | barbacoense , 1970 | Upper Albian | Barbacoas, Venezuela | , 1970 |
| 4 | bussoni , 1965 | Upper Albian | Tebaga de Kebili, South Tunisia | , 1965 |
| 5 | caneroti & , 1981 | uppermost Albian | Castellon, Spain | & , 1981 |
| Lower Albian | Maestrat Basin, Spain | et alii, 2008a | ||
| 6 | champaraense , 1956 | Middle Albian | Sihuas, northern Peru | , 1956 |
| Lower Albian | La Cuenca, Peru | et alii, 2003 | ||
| 7 | discoides , 1970 | Upper Albian | Barbacoas, Venezuela | , 1970 |
| 8 | duartei (, 1979) | Lower Albian | Corral Quemado, Peru | et alii, 2003; & , 2004 |
| 9 | duboisi (, 1989) | Upper Albian | Salazac, Gard, France | , 1989; , 2008 |
| Bakony Mts., Hungary | , 1990a, 1990b | |||
| 10 | duplicatum , 1970 | Upper Albian | Barbacoas, Venezuela | , 1970 |
| Basque-Cantabrian Basin, N Spain | , 1970; et alii, 2009 | |||
| 11 | ebrayi (, 1882) | Albian | Cosne, France | , 1928, , 1938 |
| Upper Albian | Santander, Cantabria, Spain | , 1996 | ||
| Middle Albian | France | et alii, 1996 | ||
| Perchois, Aube, France | , 1979; , 2008 | |||
| Lower Albian | Peru | et alii, 2003; & , 2004 | ||
| Maestrat Basin, Spain | et alii, 2008a | |||
| 12 | elegans , 1970 | Upper Albian | Barbacoas, Venezuela | , 1970 |
| 13 | guadaloupaeforme (, 1910) | Middle Albian | northern Peru | , 1910; et alii, 1998 |
| Cajamarca, Condebamba, northern Peru | , 1956; et alii, 1998 | |||
| Lower Albian | La Cuenca, Peru | et alii, 2003 | ||
| 14 | haasi , 1956 | Lower Albian | Celendín, northern Peru | , 1956 |
| upper Lower Albian | Peru | et alii, 2003; & , 2004 | ||
| 15 | icaunensis (, 1853) | Albian | Ravennes, Yonne, France | , 1853; , 1978; , 2008 |
| 16 | mahmoudi (, 1953) | lowermost Albian | Mellégue, Algeria | , 1953; & , 2004 |
| England | , 1978 | |||
| 17 | nodosum (, 1940b) | Upper Albian | Glen Rose, Texas | , 1940b |
| 18 | pernodosum (, 1910) | Lower Albian | Cajabamba, northern Peru | , 1910; et alii, 2003; & , 2004 |
| Middle Albian | northern Peru | , 1956; et alii, 1998 | ||
| La Cuenca, Peru | et alii, 2003 | |||
| 19 | roemeri (, 1893) | Cenomanian | Texas | , 1905 |
| Upper Albian | Glen Rose, Texas | , 1940b | ||
| Albian | Texas | , 1903; et alii, 1998a | ||
| 20 | tetranodosum (, 1925) | Middle Albian | Celendín, Cajamarca, northern Peru | , 1956 |
| uppermost Lower Albian | Cajamarca, northern Peru | et alii, 1998 | ||
| 21 | trinitense (, 1940b) | Upper Albian | Fort Worth, Texas | , 1940b |
| 22 | zagrosiense , 1981 | ? Lower Cenomanian | Kazhdumi, southern Iran | , 1981 |
| 23 | ziczag (, 1952) | Middle Albian | Uchupata, northern Peru | , 1956; et alii, 1998; & , 2004 |
| Platiknemiceras , 1954 | uppermost Aptian-Lower Cenomanian | Tethyan | ||
| 1 | bassei , 1954 | Lower-Middle Albian | Spain | , 1961 |
| Lower Albian | et alii, 1996 | |||
| Maestrat Basin, Cataluna, Spain | et alii, 1994; et alii, 2008a | |||
| uppermost Aptian | SW Iran | , 2010 | ||
| 2 | caseyi et alii, 1980 | Lower Albian | Kyushu, Japan | et alii, 1980 |
| 3 | colombiana , 1979 | upper L. Albian-lower M. Albian | Colombia | , 1979, 1981; , 1999 |
| 4 | deserti (, 1955) | Upper Albian | Utrillas Formation, SE Spain | et alii, 1993 |
| Lower Albian | Moghara, Egypt | , 1955; & , 1955; , 1961 | ||
| Lurestan, Iran | , 2007 | |||
| 5 | flexuosum et alii, 1998a | Lower-Middle Albian | Texas, USA | et alii, 1998a |
| upper Lower Albian | Peru | & , 2004 | ||
| 6 | gracile (, 1916) | Albian | Mderedj, Lebanon | , 1940; , 1961 |
| Upper Albian | Utrillas Formation, Spain | et alii, 1993 | ||
| Oued Zitoun, North Tunisia | & , 1986 | |||
| Gafsa, Central Tunisia | & , 2008 | |||
| Middle Albian | Tebaga, central Tunisia | , 1965; et alii, 1985 | ||
| Dead Sea, Israel | , 2002 | |||
| Lower Albian | Moghara, northern Sinai, Egypt | , 1928; & , 1955; & , 1999; , 2001; , 2008 | ||
| 7 | hachourii (, 1953) | lowermost Albian | Mellégue, Algeria | , 1953; & , 2004 |
| basal Albian | Guajira, NE Colombia | , 2002 | ||
| 8 | jullieni (, 1940) | Upper Albian | El Mizab, Meknes, Atlas, Morocco | & , 1968 |
| Albian | Khenchela, Algeria | , 1961 | ||
| 9 | sequanense , 1979 | Lower-Middle Albian | Aube, France | , 1979 |
| 10 | subcomplicatum (, 1940) | Albian | Lebanon | , 1940; , 1961 |
| 11 | valencianum & , 1980 | uppermost Albian-lowermost Cenomanian | Valencia, Spain | & , 1980; et alii, 2004 |
Table 1: Geographical and stratigraphical distribution of engonoceratid species. Type species are indicated in bold.