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Revision of some problematic Early Cretaceous ammonite species described by H. Coquand (1880)
Cretaceous Research 32 (2011) 187e193
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Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes
Revision of some problematic Early Cretaceous ammonite species described by H. Coquand (1880) Ottilia Szives a, *, Miguel Company b a b
Geological and Palaeontological Department, Hungarian Natural History Museum, 1431 Budapest, pf. 137, Hungary Departamento de Estratigrafía y Paleontología, Facultad de Ciencias, Universidad de Granada, 18002 Granada, Spain
a r t i c l e i n f o
a b s t r a c t
Article history: Received 6 April 2010 Accepted in revised form 1 December 2010 Available online 14 December 2010
The finding and analysis of the type specimens of some Barremian ammonite species described, but not depicted by Coquand (1880, Bull. de l’Acad. d’Hippone 15, 1-449), has led us to the revise their current interpretation. The purpose of the manuscript is to re-describe the three most commonly used Early Cretaceous species described by Coquand (1880, Bull. de l’Acad. d’Hippone 15, 1-449) and is to consider one species as a nomen dubium. The original type material of Holcodiscus diversecostatus, Holcodiscus metamorphicus and Heinzia heinzi shows that these species have been usually misidentified in the literature. We designate lectotypes for these species, thus amending some previous invalid designations, and fully discuss their actual status and synonymy. In addition, we conclude that the use of the nominal species “Ammonites mazuca” should be fully avoided, it is considered as a nomen dubium. The type material is apparently seems to be lost so its application is very doubtful. Ó 2010 Elsevier Ltd. All rights reserved.
Keywords: Ammonites Early Cretaceous Barremian Coquand Types Algeria
1. Introduction Henri Coquand (1813e1881) was an influential geologist of his time. Professor of Geology and Mineralogy at the universities of Besançon and Marseille, he carried out many geological and palaeontological studies in different European and North-African areas. Shortly after his death, his huge paleontological collection, mainly composed of materials from Algeria, France and Spain, was sold by his inheritors to Count Andor Semsey, the greatest patron of the Hungarian science, who immediately (in 1882) donated the collection to the Hungarian Geological Institute, where it is still held today. In his last publication, Coquand (1880) described, among many other invertebrate fossils from the Jurassic and Cretaceous of the Constantine region (northeastern Algeria), more than 30 new species of ammonites from the “Neocomian” (in reality, Barremian and Aptian) of Djebel Ouach, Duvivier (now Bouchegouf) and some other localities. Unfortunately, Coquand provided no figures and only short diagnoses of the new taxa, which made very difficult their identification. In 1886, Charles Heinz, a local amateur palaeontologist who had collected many of the fossils studied by Coquand, printed five plates with pictures of many of the species described by Coquand. Nevertheless, as the original type specimens
* Corresponding author. Tel.: þ36 208080958; fax: þ36 13382728 E-mail address: [email protected] (O. Szives). 0195-6671/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2010.12.001
had remained in the Coquand Collection, then already moved to Budapest. Heinz had to trust his memory in order to identify the specimens he figured (Sayn, 1891; Papier, 1896). This would explain the patent divergences between some of the Heinz’s figures and the original diagnoses by Coquand. Despite this and the fact that his plates were never effectively published, most of the subsequent authors have followed the interpretations of Heinz. The present work has been developed in the framework of a project led by one of us (O. Szives) and set in motion in 2005 in order to review the remaining Cretaceous ammonite material of the Coquand Collection in the Geological Museum of Hungary (GMH). The scientific importance of Coquand’s ammonite collections is well-known by the specialists, due to his 109 published, new Cretaceous ammonite species, from which approximately two dozens are in common use, besides the more-or-less dozen biostratigraphically important species. Unfortunately, the decades of wars and the political situation led to the slight scientific forgottening of the collection and most of the publications after Pervinquiére (1910) cited Coquand’s types as “lost” items. The purpose of the manuscript is to re-describe the four most commonly used Early Cretaceous species described by Coquand (1880), of which original types were found in the collection. Because of their important stratigraphic position and the commonly used, widespread misinterpretation of them, these specimens are taken out from the more than 30 new Early Cretaceous species of Coquand and described separately from the rest of the material hoping to be published soon in a monograph.
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Here we revise the usual interpretation of four Barremian species in the light of the rediscovered type material. These species are: Holcodiscus diversecostatus, Holcodiscus metamorphicus, and Heinzia heinzi. We also include “Ammonites mazuca” Coquand in the present discussion due to the recently published misleading statements on its type material that, unfortunately, seems to be lost from the Coquand Collection. 2. Material studied All the specimens described below are preserved as pyritized internal moulds corresponding to early whorls of the phragmocone. They belong to the former Coquand Collection, reposited in the Geological Museum of Hungary (GMH), and are temporarily housed at the Department of Palaeontology and Geology of the Hungarian Natural History Museum. In the descriptions, the following abbreviations are used for measured parameters: (D) for the mould diameter, (U) for the diameter of the umbilicus, (Wh) for the whorl-height and (Wb) for the whorl breadth. (Wb)/(Wh) ratio is also given. All measurements are expressed in millimetres. 3. Paleontologic descriptions Order Ammonitida Agassiz, 1847 Suborder Ammonitina Hyatt, 1889 Superfamily Perisphinctoidea Steinmann, 1890 Family Holcodiscidae Spath, 1923 Genus Holcodiscus Uhlig, 1882 Type species: Ammonites caillaudianus d’Orbigny, 1850 Holcodiscus diversecostatus (Coquand, 1880) Plate 1, Fig. 1A and B
* 1880 Ammonites diverse-costatus Coquand, p. 19 non 1886 Ammonites diverse-costatus Coquand e Heinz, pl. 1 pars 1890 Holcodiscus diverse-costatus Coq. e Nicklès, p. 26; pl. 1, figs. 21e24; pl. 2, figs. 14e19; pl. 4, fig. 1; text-figs. 29e34; non pl. 1, fig. 20a, b non 1891 Holcodiscus diverse-costatus Coquand e Sayn, p. 183; pl. 3, figs. 1aed, 2a, b non 1907 Holcodiscus diverse-costatus Coq. e Karakasch, p. 118; pl. 9, figs. 15a, b, 16 1907 Holcodiscus Nicklesi Karakasch, p. 119 non 1912 Holcodiscus diversecostatus Coq. e Joleaud, p. 123; pl. 1bis, figs. 24e25 1935 Holcodiscus diversecostatus Coq. e Tzankov, p. 82; pl. 6, figs. 8e9 non 1960 Holcodiscus diverse-costatus Coquand e Drushchits, p. 305; pl. 46, fig. 8a, b ? 1966 Holcodiscus nicklesi Karakasch e Breskovski, p. 104; pl. 10, fig. 4 non 1967 Holcodiscus diversecostatus (Coquand) e Dimitrova, p. 158; pl. 78, fig. 8 ? 1967 Holcodiscus nicklesi Karakasch e Dimitrova, p. 159; pl. 78, fig. 3 (¼Breskovski, 1966, pl. 10, fig. 4) non 1985 Holcodiscus diversecostatus diversecostatus (Coquand) e Tzankov and Breskovski, p. 34; pl. 8, figs. 23e24 ? 1985 Holcodiscus nicklesi Karakasch e Tzankov and Breskovski, p. 36; pl. 9, figs. 3e4 (¼Breskovski, 1966, pl. 10, fig. 4) non 1995 Holcodiscus diversecostatus (Coquand) e Company et al., fig. 8c non 1995a Holcodiscus diversecostatus (Coquand) e Avram, p. 22; pl. 4, fig. 22 non 1995b Holcodiscus cf. diversecostatus (Coquand) e Avram, pl. 18, fig. 30a, b
Fig. 1. (A) Holcodiscus diversecostatus (Coquand, 1880), lateral view, lectotype, GMH K8128, Duvivier, Algeria. (B) Holcodiscus diversecostatus (Coquand, 1880), apertural view, lectotype, GMH K8128, Duvivier, Algeria. (C) Holcodiscus metamorphicus (Coquand, 1880), lateral view, lectotype, GMH K8086a, Djebel Ouach, Algeria. (D) Holcodiscus metamorphicus (Coquand, 1880), apertural view, lectotype, GMH K8086a, Djebel Ouach, Algeria. (E) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K8087a, Djebel Ouach, Algeria. (F) Holcodiscus metamorphicus (Coquand, 1880), apertural view, GMH K8087a, Djebel Ouach, Algeria. (G) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K8127a, Duvivier, Algeria. (H) Holcodiscus metamorphicus (Coquand, 1880), ventral view, GMH K8127a, Duvivier, Algeria. (I) Holcodiscus cf. metamorphicus (Coquand, 1880), lateral view, GMH K8127b, Duvivier, Algeria. (J) Holcodiscus cf. metamorphicus (Coquand, 1880), ventral view, GMH K8127b, Duvivier, Algeria. (K) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K9163b, Djebel Ouach, Algeria. (L) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K9163a, Djebel Ouach, Algeria. (M) Holcodiscus metamorphicus (Coquand, 1880), ventral view, GMH K9163a, Djebel Ouach, Algeria. (N) Heinzia heinzi (Coquand, 1880), lateral view, lectotype, GMH K8029, Djebel Nador, Algeria. (O) Heinzia heinzi (Coquand, 1880), apertural view, lectotype, GMH K8029, Djebel Nador, Algeria. All specimens illustrated are magnified twice and temporarily borrowed and deposited in the Department of Palaeontology and Geology of the Hungarian Natural History Museum, but are repositored property of the Geological Museum of Hungary (GMH).
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non 2005 Metahoplites diverse-costatus (Coquand) e Vermeulen, pl. 44, fig. 3 ? 2006 Metahoplites cf. nicklesi (Karakasch) e Vasí cek, p. 427; pl. 2, figs. 2e3 } zy and non 2009 Holcodiscus ex gr. diversecostatus (Coquand) e Fo Janssen, fig. 4v
3.1. Original diagnosis “Diamètre: 0.023 m. Coquille peu épaisse; côtés peu convexes, ornés de côtes nombreuses, simples à leur point de départ, c’est-àdire de l’ombilic, se bifurquant, pour le plus grand nombre, au milieu du tour et venant se refermer vers le pourtour extérieur, où elles se terminent par un tubercule épineux, de manière à former une véritable ganse. Dos lisse, creusé peu profondément. Spire composée de tours comprimés, apparents dans l’ombilic sur le tiers de leur largeur”. 3.2. Type material According to Klein (2005, p. 134), the holotype of this species (by original designation by Sayn, 1891) would be the specimen figured by Sayn (1891, pl. 3, fig. 2). In fact, Sayn referred to this specimen as “le type”, but he probably meant no more than “typical morphology”. In any case, this specimen cannot be the type, because it did not belong to the type series (according to Sayn, this specimen was provided to him by Kilian). Moreover, the holotype of a species can only be fixed in the original publication and by the original author (art. 73.1.3 of ICZN). In consequence, a lectotype must be designated for this species. We here designate as lectotype the sole specimen attributed to this species conserved in the Coquand Collection (repository number GMH K8128). According to the original label, it comes from the Neocomian of Duvivier. This specimen fits well the original diagnosis by Coquand included above. 3.3. Dimensions Lectotype GMH K8128. D: 19; Wh: 9.5; Wb: 8.5; Wb/Wh: 0.89; U: 4.5 3.4. Description A tiny, 19 mm in diameter, specimen with moderate evolution. Whorl section is suboval, very slightly compressed, with convex sides, and the maximum whorl breadth is at midflank. The ornamentation consists of 23, slightly prorsiradiate, non-flexuous “primary ribs” that, in general, bifurcate at midflank. Small bullae are present at the bifurcation point in the first-half of the last visible whorl, but they disappear rapidly during the ontogeny. Secondary ribs join in twos to form together a prominent tuberculated clavus on the ventrolateral edge. At the preserved stage, the venter is broad, smooth, and flat or slightly concave.
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unaware of the existence of the Heinz’s plates, included two different morphologies in H. diversecostatus. He figured a unique specimen from Sierra de Fontcalent (pl. 1, fig. 20a, b) that was very similar to that figured by Heinz. But all the other specimens, coming from La Querola, that he included in this species (pl. 1, figs. 21e24; pl. 2, figs. 14e19), closely match the type conserved in the Coquand Collection. However, Karakasch (1907), following the interpretation of Heinz (1886) and Sayn (1891), separated these latter forms of Nicklès as a new species that he called H. nicklesi. This has been the generally accepted status, but the finding of the Coquand’s type material (repository number is: GMH K8128) leads to change our interpretation of H. diversecostatus, excluding of this species most of the specimens classically included in it, and due to its similar original description and figuration, considering H. nicklesi its junior synonym (see the synonymy list above).
3.6. Distribution This species has been reported, generally under the name of H. nicklesi, throughout the Mediterranean area (Algeria, Spain, France, Hungary, and Bulgaria). It seems to be restricted to the lower part of the Kotetishvilia compressissima Zone (Lower Barremian) }zy and Janssen, 2009). (Company et al., 1995; Vermeulen, 1996; Fo Holcodiscus metamorphicus (Coquand, 1880) Plate 1, Fig. 1CeM * 1880 Ammonites metamorphicus Coquand, p. 20 1886 Ammonites diverse-costatus Coquand e Heinz, pl. 1 non 1886 Ammonites metamorphicus Coquand e Heinz, pl. 1 pars 1890 Holcodiscus diverse-costatus Coq. e Nicklès, p. 26; pl. 1, fig. 20a, b; non pl. 1, figs. 21e24; pl. 2, figs. 14e19; pl. 4, fig. 1; text-figs. 29e34 1891 Holcodiscus diverse-costatus Coquand e Sayn, p. 183; pl. 3, figs. 1aed, 2a, b pars? 1891 Holcodiscus Geronimae Hermite e Sayn, p. 186; ? pl. 3, fig. 5; non pl. 3, fig. 4 ? 1907 Holcodiscus diverse-costatus Coq. e Karakasch, p. 118; pl. 9, figs. 15a, b, 16 non 1912 Holcodiscus metamorphicus Coq. e Joleaud, p. 123; pl. 1bis, figs. 18e20 1912 Holcodiscus diversecostatus Coq. e Joleaud, p. 123; pl. 1bis, figs. 24e25 1960 Holcodiscus diverse-costatus Coquand e Drushchits, p. 305; pl. 46, fig. 8a, b ? 1985 Holcodiscus diversecostatus diversecostatus (Coquand) e Tzankov and Breskovski, p. 34; pl. 8, figs. 23e24 1995 Holcodiscus diversecostatus (Coquand) e Company et al., fig. 8c 1995a Holcodiscus diversecostatus (Coquand) e Avram, p. 22; pl. 4, fig. 22 ? 1995b Holcodiscus cf. diversecostatus (Coquand) e Avram, pl. 18, fig. 30a, b 2004 Holcodiscus cf. nicklesi Karakasch e Lukeneder, pl. 2, fig. 4 2005 Metahoplites diverse-costatus (Coquand) e Vermeulen, pl. 44, fig. 3
3.5. Discussion H. diversecostatus is a species commonly cited in the literature as it can be seen above at the synonymy. Sayn (1891) and most of the subsequent authors have interpreted it according to the specimen figured by Heinz (1886). The Heinz specimen, however, is quite different from the lectotype above described as it has a finer and more flexuous ribbing. Nicklès (1890), who apparently was
3.7. Original diagnosis “Diamètre: 0.018 m. Espèce variable suivant l’âge et les individus. Jeune, elle débute par des tours arrondis sur le dos, ornés de côtes fines bi ou trifurquées, lesquelles passent de l’autre côté. Adulte, les côtes se montrent plus espacées, et, de distance en
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distance, mais sans qu’on puisse observer de bien régulier dans leur espacement, quelques-unes deviennent plus saillantes et séparent les côtes intermédiaires en faisceaux d’inégale dimension. Le dos, d’abord arrondi, se creuse progressivement, à mesure que la coquille prend de l’accroissement et présente un méplat que dominent, de chaque côte, des côtes armées d’un tubercule épineux. Spire composée de tours arrondis se recouvrant sur la moitié de leur largeur et laissant place pour un ombilic assez profond”.
3.8. Type material The Coquand Collection houses a number of pyritic nuclei referred to “metamorphicus”, but likely corresponding to more than one single species. We have chosen as lectotype one of the most representatives, to which the majority of the other specimens can be compared. It is labelled with number GMH K8086a and comes from Djebel Ouach. Once the lectotype has been designated, all the other specimens of the type series automatically become paralectotypes (art. 73.2.2 of ICZN).
by a single specimen from Djebel Ouach, with lateral tubercles. The specimen figured by Heinz (1886) corresponds to this latter tuberculate form that Sayn (1891) and Fallot and Termier (1923) considered to be the same as H. geronimae (Hermite), although this view was not shared by other authors like Kilian (1910) and Joleaud (1912). In any case, the great majority of the specimens labelled as Am. metamorphicus in the original material, including the lectotype here chosen, is very different from this tuberculate form and closely resemble the specimen figured by Heinz (1886) as Am. diversecostatus. As a consequence, most of the specimens referred to H. diversecostatus by the subsequent authors, who in the main have followed the Heinz interpretation, correspond in fact to H. metamorphicus. 3.12. Distribution This species is a common component of the ammonite assemblages of the upper part of the K. compressissima Zone and the lower part of the Moutoniceras moutonianum Zone (Lower Barremian). It has been found in Algeria, Spain, France, Austria, Rumania and Crimea.
3.9. Dimensions Lectotype. GMH K8086a. D: 17; Wh: 8.2; Wb: 7.6; Wb/Wh: 0.92; U: 3 Paralectotype 1. GMH K8087a. D: 13; Wh: 6.5; Wb: 6.2; Wb/Wh: 0.95; U: 2 Paralectotype 2. GMH K8127a. D: 13.5; Wh: 6; Wb: 6.5; Wb/Wh: 1.08; U: 2 Paralectotype 3. GMH K9163a. D: 12.5; Wh: 6; Wb: 6.5; Wb/Wh: 1.08; U: 2 3.10. Descriptions 3.10.1. Lectotype “GMH K8086a” Small, involute form with subcircular, slightly compressed whorl section with the maximum whorl breadth at midflank. The ornamentation consists of very fine, crowded and flexuous primary ribs arising from the umbilical seam. Primary ribs bi- or trifurcate at midflank and also intercalated ribs can be observed. Some of the secondary ribs join by twos at a small ventrolateral tubercle. The venter is broad, flattened and smooth at this stage. 3.10.2. Figured paralectotypes GMH K8087a. Similar to the lectotype, but with coarser ornamentation at the same diameter. GMH K8127a. Similar to GMH K8087a, but corresponding to a more advanced ontogenetic stage. Tubercles are very well developed and ribs begin to cross the venter. GMH K9163a. Similar to the lectotype, with a slightly more circular whorl section. We also figure a specimen (GMH K8127b, Plate 1, Fig. 1I and J), that we attribute to this species with certain reservations. It is similar to the strongly ornamented morphotypes but, in addition, it shows narrow, periodical constrictions adapically flanked by very strong ribs. This specimen is figured as H. cf. metamorphicus. 3.11. Discussion As we can be seen above, the original diagnosis is quite vague and could be applied to many different holcodiscids. Moreover, Coquand (1880) also included in this species a variety, represented
Family Pulchelliidae Douvillé, 1890 Genus Heinzia Sayn, 1891 Type species: Heinzia sayni Hyatt, 1903 Heinzia heinzi (Coquand, 1880) Plate 1, Fig. 1N and O * 1880 Ammonites Heinzi Coquand, p. 18 non 1886 Ammonites Heinzi Coquand e Heinz, pl. 1 non 1891 Pulchellia Heinzi Coquand in Heinz e Sayn, p. 159; pl. 2, fig. 5aec ? 1891 Pulchellia subcaicedi nov. sp. e Sayn, p. 163, pl. 2, fig. 6 non 1966 Pulchellia heinzi (Coquand) e Breskovski, p. 108; pl. 5, fig. 5 non 1967 Pulchellia heinzi (Coquand) e Dimitrova, p. 166; pl. 80, fig. 6 (¼Breskovski, 1966, pl. 5, fig. 5) 1980a Pulchellia (Heinzia) heinzi (Coq.) e Vermeulen, pl. 4, figs. 11e14 1980b Pulchellia (Heinzia) heinzi (Coq.) e Vermeulen, pl. 6, figs. 11e14 (¼Vermeulen, 1980a, pl. 4, figs. 11e14) non 1995 Coronites (Curiolites) aff. heinzi (Coquand) in Sayn e Vermeulen, pl. 2, fig. 9 non 1997 Curiolites aff. heinzi (Sayn) e Vermeulen, pl. 3, fig. 12 (¼Vermeulen, 1995, pl. 2, fig. 9) 1998a Heinzia gr. sayni Hyatt e Vermeulen, pl. 4, figs. 4e6 (¼Vermeulen, 1980a, pl. 4, figs. 12e14) non 2003 Curiolia heinzi (Sayn) e Vermeulen, p. 158; pl. 41, figs. 5 (¼Vermeulen, 1995, pl. 2, fig. 9), 9e11 2003 Heinzia sp. gr. sayni Hyatt e Vermeulen, pl. 47, fig. 4 (¼Vermeulen, 1980a, pl. 4, fig. 14 3.13. Original diagnosis “Diamètre: 0.017 m. Coquille de petit taille, un peu comprimée, ornée par tours de quatorze côtes simples quelquefois, mais plus souvent bifurquées, partant de l’ombilic, espacées, plates et épaisses, s’arrêtant au milieu du dos, où elles s’infléchissent légèrement en avant en se terminant par un tubercule obtus et tronqué; dos creusé par un sillon profond et étroit, dominé par les tubercules de côtes. Spire formée de tours convexes légèrement anguleux, apparents dans l’ombilic sur la moitié de leur largeur. Bouche subquadrangulaire”.
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3.14. Type material
3.18. Distribution
The only specimen of this species remaining in the Coquand Collection (GMH K8029) is smaller than that originally described by Coquand (1880) and comes from a different locality. Nevertheless, it fits well the original description. We here designate it as lectotype of the species. According to the original label of Coquand, it comes from Djebel Nador. Another specimen, labelled GMH K8097, was a repositored item and had been listed in 1984. It probably was the specimen described by Coquand (1880). Unfortunately all our efforts to find this specimen now have been failed. Vermeulen (2003) designated the specimen of Sayn (1890, pl. 2, fig. 5) as lectotype of C. heinzi Sayn (non Coquand), as he correctly thought that the figured specimen by Sayn did not belong to the same species as those described by Coquand (1880). Therefore, the designation of the lectotype by Vermeulen for C. heinzi Sayn (non Coquand) is not relevant for this paper.
All the specimens that can be undoubtedly attributed to this species (see synonymy list) come from the Barremian of Algeria. Its exact stratigraphic position is not known, but its ornamentation is similar to that of other pulchelliids from the M. moutonianum Zone. Undetermined Family
3.15. Dimensions Lectotype GMH K8029. D: 11; Wh: 5; Wb: 4; Wb/Wh:0.85; U: 2
3.16. Description Tiny, pyritic, rather involute and compressed nucleus of 11 mm diameter. The whorl section is subrectangular, with slightly convex sides and the maximum breadth at midflank. The ornamentation consists of broad, flat ribs that arise, mostly in pairs, from an umbilical swell and develop a strong, truncated tubercle on the outer lateral region. The ventral region is smooth, narrow and concave.
3.17. Discussion Heinz (1886) figured as belonging to this species a specimen that did not correspond to the Coquand description. In the Heinz specimen most of the ribs are simple and rigid, and the ventrolateral tubercles are nodular. Sayn (1891), even recognizing these differences, followed the Heinz interpretation. Furthermore, he described a new species, P. subcaicedi, which showed strong agreement with the Coquand diagnosis of “Ammonites heinzi”. On the other hand, Vermeulen (1980a, b) attributed two specimens to Pulchellia (Heinzia) heinzi from Djebel Ouach that closely match the original description and the specimen held in the Coquand Collection. More recently, however, Vermeulen (1998a, 2003) and Vermeulen and Klein (2006) have related these specimens to H. sayni Hyatt, despite the fact this latter species has grooved ventrolateral tubercles. Meanwhile, Vermeulen (1995) defined a new genus, Curiolites, for the species “Ammonites heinzi” Coquand (1880). Shortly afterwards, however, the same author (Vermeulen, 1997, p. 103) stated that the type species of Curiolites was C. heinzi (Sayn) non Coquand (1880), since he considered that the Coquand species (i. e. heinzi) could be different and should be abandoned. This change is, evidently, against the ICZN rules, as is also the subsequent replacement (Vermeulen, 1999), without any apparent reason, of the name Curiolites by Curiolia. In our opinion, the Coquand species is perfectly valid and finds its place in the genus Heinzia, near other species, like Heinzia caicedi (Karsten, 1858) and Heinzia hispanica Hyatt, 1903 with flat, rhomboidal tubercles.
Undetermined Genus Ammonites mazuca Coquand, 1880 * 1880 Ammonites Mazuca Coquand, p. 368
3.19. Original diagnosis “Diamètre: 0.015 m. Coquille discoïdale, très-comprimée, tranchante et entière à son pourtour, marquée en travers de très légères côtes, plates, flexueuses, distantes, presque sans relief, qui partent du pourtour de l’ombilic, infléchies d’avant en arrière, disparaissant près de la carène. Spire embrassante. Ombilic à peine indiqué. Tours très comprimés. Bouche en fer de lance, très-comprimée, anguleuse en avant, fortement échancrée en arrière”.
3.20. Type Our efforts to find the type material of this species in the collections of the Hungarian Geological Institute have been in vain. Prof. W. J. Kennedy (Oxford University) kindly provided us a list from 1984 of the Cretaceous ammonite specimens of the Coquand Collection. In 1984 he visited the Hungarian Geological Institute for the purpose of research on the Coquand Collection, and the former }-Czabalay gave him the document containing the curator, L. Benko specimens existing at that moment. In the copy, a “Saynella mazuca Coquand” from Djebel Ouach, with the repository number K8079 is clearly indicated. Since then, the museum collections were rearranged, and the Coquand material was temporarily moved to countryside store-houses and then back again. Many specimens were lost in the meantime, and this seems to be also the case of the “Ammonites mazuca”. Vermeulen (1996, 1998b) mentioned that the type material of this species was conserved in the Coquand Collection at Budapest, but without giving any further information. More recently, however, Vermeulen (2003) has designated as lectotype a specimen from his own collection, which contravenes the Article 74 of the Code.
3.21. Discussion After its creation, this species name had never been used until Vermeulen (1996) considered it to be a senior synonym of a wellknown pulchelliid from the lowermost Barremian, Psilotissotia chalmasi (Nicklès, 1890). Since then, Psilotissotia mazuca has been repeatedly used by our colleague to characterize a unit of his successive biostratigraphic schemes for the Barremian of SE France (Vermeulen, 1996, 1998a, 2003, 2005). Nevertheless, we think that there are strong reasons to regard this species as a nomen dubium. Firstly, as stated above, its type material seems to be definitively lost. Secondly, the original description is quite imprecise and could correspond not only to a Psilotissotia, but also to different species of Arnaudiella, Torcapella, Barremites or the Aptian Pseudosaynella or Aconeceras. And lastly, no other of the ammonite species reported from Djebel Ouach (Coquand, 1880; Sayn, 1890, 1891, 1896; Joleaud, 1901, 1912;
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Busnardo, 1957; Vermeulen and Lahondère, 2008) can be exclusively attributed to the lowermost Barremian, and most of them are characteristic of the middle and upper parts of the stage (K. compressissima Zone or higher stratigraphic levels). Under these circumstances, the synonymy between “Ammonites mazuca” and P. chalmasi cannot be verified, so the use of the nominal species proposed by Coquand should be avoided.
4. Summary and conclusions Coquand (1880) established several new species of Barremian ammonites with only short descriptions and no figures, which rendered their identification difficult and controversial. The analysis, for the first time after its original description, of the Coquand’s type material, held in the Hungarian Geological Institute, has enabled us to designate the lectotypes and revise the current conceptions of some of these species. H. diversecostatus, H. metamorphicus and H. heinzi have been misidentified by most of the authors, who have adopted the interpretation by Heinz (1886) of these species. H. diversecostatus (Coquand) is here considered to be a senior synonym of H. nicklesi (Karakasch), whereas most of the specimens attributed by the authors to H. diversecostatus correspond, in fact, to true H metamorphicus (Coquand). In turn, H. heinzi (Coquand) has to be regarded as a different species from the forms figured by Heinz (1886) and Sayn (1891) under this name. Here we also designate lectotypes for H. diversecostatus, H. metamorphicus and Hz. heinzi. The type material of “Ammonites mazuca” has not been found. This and the uncertainty on its stratigraphic position lead us to consider this species name as a nomen dubium, whose application should be fully avoided.
Acknowledgements This paper is a contribution to Project HH 2008-0014 (Hungarian-Spanish Cooperation Programme). The research of M. Company has been also co-financed by Project CGL 2008-00533 (Spanish Ministry of Science and Technology) and Research Group RNM-178 (Junta de Andalucía). The scientific research of O. Szives on the Coquand Collection was supported by the Hungarian OTKA K62063 research project and the TéT ES 28/2008 HungarianSpanish Cooperation Programme. The authors are also grateful to Prof. W. J. Kennedy (Oxford University Museum) for providing them his notes on the Coquand Collection.
References Agassiz, J.L.R., 1847. Nomenclator Zoologicus. Fasciculus XII. Indicem Universalem. Nomenclatoris Zoologici Index Universalis, continens nomina systematica classium, ordinum, familiarum et generum animalium omnium, tam vivetium quam fossilium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plantarum, nec non variis adnotationibus et emendationibus. Jent et Gassmann VII, 393. Avram, E., 1995a. Representatives of the Family Holcodiscidae Spath, 1924 (Ammonitina) in Rumania. Memorie Descritive della Carta Geologica d’Italia 51, 11e45. Avram, E., 1995b. Lower Cretaceous (Valanginian-Early Aptian) ammonite succession in the Svinit¸a region (SW Rumania). Géologie Alpine, Mémoire Hors Série 20 (1994), 113e167. Breskovski, S., 1966. Biostratigraphie du Barrémien au sud du village de Brestak, dans la région de Varna. Travaux sur la Géologie de Bulgarie, Série Paléontologie 8, 71e121. 10 pls. (In Bulgarian, French summary). Busnardo, R., 1957. Étude stratigraphique des marnes à ammonoïdés de la région de Constantine. Bulletin du Service de la Carte Géologique de l’Algérie, Nouvelle Série 13, 7e65. Company, M., Sandoval, J., Tavera, J.M., 1995. Lower Barremian ammonite biostratigraphy in the Subbetic Domain (Betic Cordillera, southern Spain). Cretaceous Research 16, 243e256.
Coquand, H., 1880. Études supplémentaires sur la paléontologie algérienne faisant suite à la description géologique et paléontologique de la région sud de la province de Constantine. Bulletin de l’Académie d’Hippone 15, 1e449. Dimitrova, N., 1967. Les fossiles de Bulgarie. IV. Crétacé inférieur. Cephalopoda (Nautiloidea et Ammonoidea). Académie Bulgare des Sciences, Sofia, 424 pp. (In Bulgarian, French summary). d’Orbigny, A., 1850. Notes sur quelques nouvelles espéces remarquables d’Ammonites des étages Néocomien et Aptien de la France. Journal de Conchyliologie, Paris I, 196e201. pl. 8. Drushchits, V.V., 1960. Ammonites. Part I. In: Drushchits, V.V., Kudriavtsev, M.P. (Eds.), Atlas of the Lower Cretaceous fauna of the northern Caucasus and Crimea. Gostoptekhizdat, Moscow, pp. 249e308. 47 pls. (In Russian). Fallot, P., Termier, H., 1923. Ammonites nouvelles des Îles Baléares. Trabajos del Museo Nacional de Ciencias Naturales, Serie Geológica 32, 1e32. 6 pls. }zy, I., Janssen, N.M.M., 2009. Integrated Lower Cretaceous biostratigraphy of the Fo Bersek Quarry, Gerecse Mountains, Transdanubian Range, Hungary. Cretaceous Research 30, 78e92. Heinz, C., 1886. Fossiles décrits par H. Coquand dans le Bulletin de l’Académie d’Hippone 15. 5 pls. (Unpublished). Hyatt, A., 1889. Cephalopoda. p. 502e592, fig. 1049e1235. In: Zittel, K.A. (Ed.), Textbook of Palaeontology. Macmillan, London & New York. Joleaud, A., 1901. Contribution à l’étude de l’Infracretacé à faciès vaseux pélagique en Algérie et en Tunisie. Bulletin de la Société Géologique de France, Quatrième Série 1, 113e146. Joleaud, L., 1912. Étude géologique de la Chaîne Numidique et les monts de Constantine (1911). Montane-Sicardi et Valentin, Montpellier, 437. 6 pls. Karakasch, N.I., 1907. Le Crétacé inférieur de la Crimée et sa faune. Travaux de la Société Impériale des Naturalistes de St.-Pétersbourg 32 (5), 1e482. 28 pls. (In Russian, French summary). Karsten, 1858. Über die geognostischen Verhältnisse des Westlichen Colombien des heutigen Republiken Neu Granada und Ecuador. Amtlicher Bericht der Versammlung Deutscher Naturforscher und Aertze 32 (1856), 80e117. Kilian, W., 1910. II Teil: Das Mesozoicum. 3 Band: Kreide. Erste Abteilung: Unterkreide (Palaeocretacicum). Lieferung 2: Das bathyale Palaeocretacicum im südöstlichen Frankreich. Valendis-Stufe; Hauterive-Stufe; Barrême-Stufe; Apt-Stufe. In: Frech, F. (Ed.), Lethaea Geognostica. Schweizerbart, Stuttgart, pp. 169e288. 8 pls. Klein, J., 2005. Pars 139. Lower Cretaceous Ammonites I. Perisphinctaceae 1: Himalayitidae, Olcostephanidae, Holcodiscidae, Neocomitidae, Oosterellidae. In: Riegraf, W. (Ed.), Fossilium Catalogus I: Animalia. Backhuys Publishers, Leiden, X þ 484 pp. Lukeneder, A., 2004. Stratigrafische Erkenntnisse aus einem neuen Vorkommen von Unterkreide-Ammonoideen in der Losensteiner Mulde (Ternberger Decke, Nördliche Kalkalpen). Jahrbuch der Geologischen Bundesanstalt 144, 173e189. Nicklès, R., 1890. Contributions à la paléontologie du sud-est de l’Espagne. Terrain Crétacé. I. Néocomien. Mémoires de la Société Géologique de France, Paléontologie 4, 1e30. pls. 1e4. Papier, A., 1896. Notice sur les recherches et découvertes paléontologiques de Charles Heinz. Bulletin de l’Académie d’Hippone 28, 98e127. Pervinquiére, L., 1910. Sur quelques ammonites du crétacé algérien. Mémoires de la Societé Géologique de France, Paleontologie 17 (2e3), 86. figs. 1e38, pls. 1e7. Sayn, G., 1890. Sur la faune d’ammonites pyriteuses barrémiennes du Djebel Ouach, province de Constantine. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences 110, 1381e1382. Sayn, G., 1891. Description des ammonitides du Barrémien du Djebel Ouach, près Constantine. Annales de la Société d’Agriculture, Histoire Naturelle et Arts Utiles de Lyon, Sixième Série 3 (1890), 135e208. 3 pls. Sayn, G., 1896. Note sur le Néocomien du Djebel Ouach. Bulletin de la Société Géologique de France, Troisième Série 24, 1162e1167. Spath, L.F., 1923. A Monograph of the Ammonoidea of the Gault. Monographs of the Palaeontographical Society, London, Part 1. pp. 1e72., fig. 1e14., pl. 1e4. Steinmann, G., 1890. Cephalopoda. In: Steinmann, G., Döderlein, L. (Eds.), Elemente der Paläontologie. W. Engelmann, Leipzig, pp. 344e475. Tzankov, V., Breskovski, S., 1985. Ammonites des familles Holcodiscidae Spath, 1924 et Astieridiscidae Tzankov & Breskovski, 1982. II. Description paléontologique. Geologia Balcanica 15 (5), 3e52. 11 pls. Tzankov, V., 1935. Notes sur le genre Holcodiscus. Annuaire de l’Université de Sofia 31 (3), 57e111 (In Bulgarian, French summary). Uhlig, V., 1882. Die Cephalopoda der Wernsdorfer Schichten. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Wien, Mathematisch e Naturwissenschaftliche Klasse 46, 127e290, pp. 1e166, pl. 1e32. Vasí cek, Z., 2006. A remarkable assemblage of Early Barremian ammonites in the Central Western Carpathians (Butkov Quarry, Slovakia). Acta Geologica Polonica 56, 421e440. Vermeulen, J., Klein, J., 2006. Pars 141. Lower Cretaceous Ammonites II. Endemocerataceae: Pulchelliidae. In: Riegraf, W. (Ed.), Fossilium Catalogus I: Animalia. Backhuys Publishers, Leiden, pp. 187e278. Vermeulen, J., Lahondère, J.C., 2008. Sur quelques espèces d’ammonites du Barrémien ultra-tellien de la région de Constantine, Algérie. I. Les Pulchelliidae (Ammonitina). Annales du Museum d’Histoire Naturelle de Nice 28, 67e111. Vermeulen, J., 1980a. Étude de la Famille des Pulcheliidae. Révision de trois espèces types du Barrémien du Sud-Est de la France. Thèse Doctorat de Specialité, Université de Nice, 92 pp., 4 pls. (Unpublished). Vermeulen, J., 1980b. Biozonation homophylétique du Barrémien du Clos de Barral (Var). In: Thomel, G. (Ed.), Ammonites. Serre, Nice, pp. 181e182. pls. 5e6.
O. Szives, M. Company / Cretaceous Research 32 (2011) 187e193 Vermeulen, J., 1995. Coronites (Coronites) darsi, nouvelle espèce de la famille des Pulchellidés (Ammonoidea). Riviéra Scientifique, 57e66 (1994). Vermeulen, J., 1996. Nouvelle biozonation du Barrémien basée sur la famille des Pulchelliidae (Ammonoidea). Géologie Alpine 71 (1995), 199e211. Vermeulen, J., 1997. Origine, classification et évolution des Pulchelliinae (Douvillé) 1911 emend. Vermeulen, 1995 (Pulchelliidae, Endemocerataceae, Ammonoidea). Géologie Alpine 72 (1996), 101e115. Vermeulen, J., 1998a. Biohorizons ammonitiques dans le Barrémien du Sud-Est de la France (de la zone à Hugii à la zone à Sartousiana). Géologie Alpine 73 (1997), 99e117.
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Vermeulen, J., 1998b. Sur quelques espèces, nouvelles ou mal connues, de la famille des Pulchelliidae (Endemocerataceae, Ammonoidea). Géologie Alpine 73 (1997), 77e97. Vermeulen, J., 1999. Sur quelques espèces de Pulchellidae (Ammonoidea, Ammonitina) de l’Hauterivien et du Barrémien du Sud-Est de la France. Annales du Muséum d’Histoire Naturelle de Nice 13 (1998), 441e477. Vermeulen, J., 2003. Étude stratigraphique et paléontologique de la famille des Pulchelliidae (Ammonoidea, Ammonitina, Endemocerataceae). Géologie Alpine, Mémoire Hors Série 42 (2002), 1e333. Vermeulen, J., 2005. Boundaries, ammonite fauna and main subdivisions of the stratotype of the Barremian. Géologie Alpine, Série “Colloques et Excursions” 7, 147e173.
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Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes
Revision of some problematic Early Cretaceous ammonite species described by H. Coquand (1880) Ottilia Szives a, *, Miguel Company b a b
Geological and Palaeontological Department, Hungarian Natural History Museum, 1431 Budapest, pf. 137, Hungary Departamento de Estratigrafía y Paleontología, Facultad de Ciencias, Universidad de Granada, 18002 Granada, Spain
a r t i c l e i n f o
a b s t r a c t
Article history: Received 6 April 2010 Accepted in revised form 1 December 2010 Available online 14 December 2010
The finding and analysis of the type specimens of some Barremian ammonite species described, but not depicted by Coquand (1880, Bull. de l’Acad. d’Hippone 15, 1-449), has led us to the revise their current interpretation. The purpose of the manuscript is to re-describe the three most commonly used Early Cretaceous species described by Coquand (1880, Bull. de l’Acad. d’Hippone 15, 1-449) and is to consider one species as a nomen dubium. The original type material of Holcodiscus diversecostatus, Holcodiscus metamorphicus and Heinzia heinzi shows that these species have been usually misidentified in the literature. We designate lectotypes for these species, thus amending some previous invalid designations, and fully discuss their actual status and synonymy. In addition, we conclude that the use of the nominal species “Ammonites mazuca” should be fully avoided, it is considered as a nomen dubium. The type material is apparently seems to be lost so its application is very doubtful. Ó 2010 Elsevier Ltd. All rights reserved.
Keywords: Ammonites Early Cretaceous Barremian Coquand Types Algeria
1. Introduction Henri Coquand (1813e1881) was an influential geologist of his time. Professor of Geology and Mineralogy at the universities of Besançon and Marseille, he carried out many geological and palaeontological studies in different European and North-African areas. Shortly after his death, his huge paleontological collection, mainly composed of materials from Algeria, France and Spain, was sold by his inheritors to Count Andor Semsey, the greatest patron of the Hungarian science, who immediately (in 1882) donated the collection to the Hungarian Geological Institute, where it is still held today. In his last publication, Coquand (1880) described, among many other invertebrate fossils from the Jurassic and Cretaceous of the Constantine region (northeastern Algeria), more than 30 new species of ammonites from the “Neocomian” (in reality, Barremian and Aptian) of Djebel Ouach, Duvivier (now Bouchegouf) and some other localities. Unfortunately, Coquand provided no figures and only short diagnoses of the new taxa, which made very difficult their identification. In 1886, Charles Heinz, a local amateur palaeontologist who had collected many of the fossils studied by Coquand, printed five plates with pictures of many of the species described by Coquand. Nevertheless, as the original type specimens
* Corresponding author. Tel.: þ36 208080958; fax: þ36 13382728 E-mail address: [email protected] (O. Szives). 0195-6671/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2010.12.001
had remained in the Coquand Collection, then already moved to Budapest. Heinz had to trust his memory in order to identify the specimens he figured (Sayn, 1891; Papier, 1896). This would explain the patent divergences between some of the Heinz’s figures and the original diagnoses by Coquand. Despite this and the fact that his plates were never effectively published, most of the subsequent authors have followed the interpretations of Heinz. The present work has been developed in the framework of a project led by one of us (O. Szives) and set in motion in 2005 in order to review the remaining Cretaceous ammonite material of the Coquand Collection in the Geological Museum of Hungary (GMH). The scientific importance of Coquand’s ammonite collections is well-known by the specialists, due to his 109 published, new Cretaceous ammonite species, from which approximately two dozens are in common use, besides the more-or-less dozen biostratigraphically important species. Unfortunately, the decades of wars and the political situation led to the slight scientific forgottening of the collection and most of the publications after Pervinquiére (1910) cited Coquand’s types as “lost” items. The purpose of the manuscript is to re-describe the four most commonly used Early Cretaceous species described by Coquand (1880), of which original types were found in the collection. Because of their important stratigraphic position and the commonly used, widespread misinterpretation of them, these specimens are taken out from the more than 30 new Early Cretaceous species of Coquand and described separately from the rest of the material hoping to be published soon in a monograph.
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Here we revise the usual interpretation of four Barremian species in the light of the rediscovered type material. These species are: Holcodiscus diversecostatus, Holcodiscus metamorphicus, and Heinzia heinzi. We also include “Ammonites mazuca” Coquand in the present discussion due to the recently published misleading statements on its type material that, unfortunately, seems to be lost from the Coquand Collection. 2. Material studied All the specimens described below are preserved as pyritized internal moulds corresponding to early whorls of the phragmocone. They belong to the former Coquand Collection, reposited in the Geological Museum of Hungary (GMH), and are temporarily housed at the Department of Palaeontology and Geology of the Hungarian Natural History Museum. In the descriptions, the following abbreviations are used for measured parameters: (D) for the mould diameter, (U) for the diameter of the umbilicus, (Wh) for the whorl-height and (Wb) for the whorl breadth. (Wb)/(Wh) ratio is also given. All measurements are expressed in millimetres. 3. Paleontologic descriptions Order Ammonitida Agassiz, 1847 Suborder Ammonitina Hyatt, 1889 Superfamily Perisphinctoidea Steinmann, 1890 Family Holcodiscidae Spath, 1923 Genus Holcodiscus Uhlig, 1882 Type species: Ammonites caillaudianus d’Orbigny, 1850 Holcodiscus diversecostatus (Coquand, 1880) Plate 1, Fig. 1A and B
* 1880 Ammonites diverse-costatus Coquand, p. 19 non 1886 Ammonites diverse-costatus Coquand e Heinz, pl. 1 pars 1890 Holcodiscus diverse-costatus Coq. e Nicklès, p. 26; pl. 1, figs. 21e24; pl. 2, figs. 14e19; pl. 4, fig. 1; text-figs. 29e34; non pl. 1, fig. 20a, b non 1891 Holcodiscus diverse-costatus Coquand e Sayn, p. 183; pl. 3, figs. 1aed, 2a, b non 1907 Holcodiscus diverse-costatus Coq. e Karakasch, p. 118; pl. 9, figs. 15a, b, 16 1907 Holcodiscus Nicklesi Karakasch, p. 119 non 1912 Holcodiscus diversecostatus Coq. e Joleaud, p. 123; pl. 1bis, figs. 24e25 1935 Holcodiscus diversecostatus Coq. e Tzankov, p. 82; pl. 6, figs. 8e9 non 1960 Holcodiscus diverse-costatus Coquand e Drushchits, p. 305; pl. 46, fig. 8a, b ? 1966 Holcodiscus nicklesi Karakasch e Breskovski, p. 104; pl. 10, fig. 4 non 1967 Holcodiscus diversecostatus (Coquand) e Dimitrova, p. 158; pl. 78, fig. 8 ? 1967 Holcodiscus nicklesi Karakasch e Dimitrova, p. 159; pl. 78, fig. 3 (¼Breskovski, 1966, pl. 10, fig. 4) non 1985 Holcodiscus diversecostatus diversecostatus (Coquand) e Tzankov and Breskovski, p. 34; pl. 8, figs. 23e24 ? 1985 Holcodiscus nicklesi Karakasch e Tzankov and Breskovski, p. 36; pl. 9, figs. 3e4 (¼Breskovski, 1966, pl. 10, fig. 4) non 1995 Holcodiscus diversecostatus (Coquand) e Company et al., fig. 8c non 1995a Holcodiscus diversecostatus (Coquand) e Avram, p. 22; pl. 4, fig. 22 non 1995b Holcodiscus cf. diversecostatus (Coquand) e Avram, pl. 18, fig. 30a, b
Fig. 1. (A) Holcodiscus diversecostatus (Coquand, 1880), lateral view, lectotype, GMH K8128, Duvivier, Algeria. (B) Holcodiscus diversecostatus (Coquand, 1880), apertural view, lectotype, GMH K8128, Duvivier, Algeria. (C) Holcodiscus metamorphicus (Coquand, 1880), lateral view, lectotype, GMH K8086a, Djebel Ouach, Algeria. (D) Holcodiscus metamorphicus (Coquand, 1880), apertural view, lectotype, GMH K8086a, Djebel Ouach, Algeria. (E) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K8087a, Djebel Ouach, Algeria. (F) Holcodiscus metamorphicus (Coquand, 1880), apertural view, GMH K8087a, Djebel Ouach, Algeria. (G) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K8127a, Duvivier, Algeria. (H) Holcodiscus metamorphicus (Coquand, 1880), ventral view, GMH K8127a, Duvivier, Algeria. (I) Holcodiscus cf. metamorphicus (Coquand, 1880), lateral view, GMH K8127b, Duvivier, Algeria. (J) Holcodiscus cf. metamorphicus (Coquand, 1880), ventral view, GMH K8127b, Duvivier, Algeria. (K) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K9163b, Djebel Ouach, Algeria. (L) Holcodiscus metamorphicus (Coquand, 1880), lateral view, GMH K9163a, Djebel Ouach, Algeria. (M) Holcodiscus metamorphicus (Coquand, 1880), ventral view, GMH K9163a, Djebel Ouach, Algeria. (N) Heinzia heinzi (Coquand, 1880), lateral view, lectotype, GMH K8029, Djebel Nador, Algeria. (O) Heinzia heinzi (Coquand, 1880), apertural view, lectotype, GMH K8029, Djebel Nador, Algeria. All specimens illustrated are magnified twice and temporarily borrowed and deposited in the Department of Palaeontology and Geology of the Hungarian Natural History Museum, but are repositored property of the Geological Museum of Hungary (GMH).
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non 2005 Metahoplites diverse-costatus (Coquand) e Vermeulen, pl. 44, fig. 3 ? 2006 Metahoplites cf. nicklesi (Karakasch) e Vasí cek, p. 427; pl. 2, figs. 2e3 } zy and non 2009 Holcodiscus ex gr. diversecostatus (Coquand) e Fo Janssen, fig. 4v
3.1. Original diagnosis “Diamètre: 0.023 m. Coquille peu épaisse; côtés peu convexes, ornés de côtes nombreuses, simples à leur point de départ, c’est-àdire de l’ombilic, se bifurquant, pour le plus grand nombre, au milieu du tour et venant se refermer vers le pourtour extérieur, où elles se terminent par un tubercule épineux, de manière à former une véritable ganse. Dos lisse, creusé peu profondément. Spire composée de tours comprimés, apparents dans l’ombilic sur le tiers de leur largeur”. 3.2. Type material According to Klein (2005, p. 134), the holotype of this species (by original designation by Sayn, 1891) would be the specimen figured by Sayn (1891, pl. 3, fig. 2). In fact, Sayn referred to this specimen as “le type”, but he probably meant no more than “typical morphology”. In any case, this specimen cannot be the type, because it did not belong to the type series (according to Sayn, this specimen was provided to him by Kilian). Moreover, the holotype of a species can only be fixed in the original publication and by the original author (art. 73.1.3 of ICZN). In consequence, a lectotype must be designated for this species. We here designate as lectotype the sole specimen attributed to this species conserved in the Coquand Collection (repository number GMH K8128). According to the original label, it comes from the Neocomian of Duvivier. This specimen fits well the original diagnosis by Coquand included above. 3.3. Dimensions Lectotype GMH K8128. D: 19; Wh: 9.5; Wb: 8.5; Wb/Wh: 0.89; U: 4.5 3.4. Description A tiny, 19 mm in diameter, specimen with moderate evolution. Whorl section is suboval, very slightly compressed, with convex sides, and the maximum whorl breadth is at midflank. The ornamentation consists of 23, slightly prorsiradiate, non-flexuous “primary ribs” that, in general, bifurcate at midflank. Small bullae are present at the bifurcation point in the first-half of the last visible whorl, but they disappear rapidly during the ontogeny. Secondary ribs join in twos to form together a prominent tuberculated clavus on the ventrolateral edge. At the preserved stage, the venter is broad, smooth, and flat or slightly concave.
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unaware of the existence of the Heinz’s plates, included two different morphologies in H. diversecostatus. He figured a unique specimen from Sierra de Fontcalent (pl. 1, fig. 20a, b) that was very similar to that figured by Heinz. But all the other specimens, coming from La Querola, that he included in this species (pl. 1, figs. 21e24; pl. 2, figs. 14e19), closely match the type conserved in the Coquand Collection. However, Karakasch (1907), following the interpretation of Heinz (1886) and Sayn (1891), separated these latter forms of Nicklès as a new species that he called H. nicklesi. This has been the generally accepted status, but the finding of the Coquand’s type material (repository number is: GMH K8128) leads to change our interpretation of H. diversecostatus, excluding of this species most of the specimens classically included in it, and due to its similar original description and figuration, considering H. nicklesi its junior synonym (see the synonymy list above).
3.6. Distribution This species has been reported, generally under the name of H. nicklesi, throughout the Mediterranean area (Algeria, Spain, France, Hungary, and Bulgaria). It seems to be restricted to the lower part of the Kotetishvilia compressissima Zone (Lower Barremian) }zy and Janssen, 2009). (Company et al., 1995; Vermeulen, 1996; Fo Holcodiscus metamorphicus (Coquand, 1880) Plate 1, Fig. 1CeM * 1880 Ammonites metamorphicus Coquand, p. 20 1886 Ammonites diverse-costatus Coquand e Heinz, pl. 1 non 1886 Ammonites metamorphicus Coquand e Heinz, pl. 1 pars 1890 Holcodiscus diverse-costatus Coq. e Nicklès, p. 26; pl. 1, fig. 20a, b; non pl. 1, figs. 21e24; pl. 2, figs. 14e19; pl. 4, fig. 1; text-figs. 29e34 1891 Holcodiscus diverse-costatus Coquand e Sayn, p. 183; pl. 3, figs. 1aed, 2a, b pars? 1891 Holcodiscus Geronimae Hermite e Sayn, p. 186; ? pl. 3, fig. 5; non pl. 3, fig. 4 ? 1907 Holcodiscus diverse-costatus Coq. e Karakasch, p. 118; pl. 9, figs. 15a, b, 16 non 1912 Holcodiscus metamorphicus Coq. e Joleaud, p. 123; pl. 1bis, figs. 18e20 1912 Holcodiscus diversecostatus Coq. e Joleaud, p. 123; pl. 1bis, figs. 24e25 1960 Holcodiscus diverse-costatus Coquand e Drushchits, p. 305; pl. 46, fig. 8a, b ? 1985 Holcodiscus diversecostatus diversecostatus (Coquand) e Tzankov and Breskovski, p. 34; pl. 8, figs. 23e24 1995 Holcodiscus diversecostatus (Coquand) e Company et al., fig. 8c 1995a Holcodiscus diversecostatus (Coquand) e Avram, p. 22; pl. 4, fig. 22 ? 1995b Holcodiscus cf. diversecostatus (Coquand) e Avram, pl. 18, fig. 30a, b 2004 Holcodiscus cf. nicklesi Karakasch e Lukeneder, pl. 2, fig. 4 2005 Metahoplites diverse-costatus (Coquand) e Vermeulen, pl. 44, fig. 3
3.5. Discussion H. diversecostatus is a species commonly cited in the literature as it can be seen above at the synonymy. Sayn (1891) and most of the subsequent authors have interpreted it according to the specimen figured by Heinz (1886). The Heinz specimen, however, is quite different from the lectotype above described as it has a finer and more flexuous ribbing. Nicklès (1890), who apparently was
3.7. Original diagnosis “Diamètre: 0.018 m. Espèce variable suivant l’âge et les individus. Jeune, elle débute par des tours arrondis sur le dos, ornés de côtes fines bi ou trifurquées, lesquelles passent de l’autre côté. Adulte, les côtes se montrent plus espacées, et, de distance en
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distance, mais sans qu’on puisse observer de bien régulier dans leur espacement, quelques-unes deviennent plus saillantes et séparent les côtes intermédiaires en faisceaux d’inégale dimension. Le dos, d’abord arrondi, se creuse progressivement, à mesure que la coquille prend de l’accroissement et présente un méplat que dominent, de chaque côte, des côtes armées d’un tubercule épineux. Spire composée de tours arrondis se recouvrant sur la moitié de leur largeur et laissant place pour un ombilic assez profond”.
3.8. Type material The Coquand Collection houses a number of pyritic nuclei referred to “metamorphicus”, but likely corresponding to more than one single species. We have chosen as lectotype one of the most representatives, to which the majority of the other specimens can be compared. It is labelled with number GMH K8086a and comes from Djebel Ouach. Once the lectotype has been designated, all the other specimens of the type series automatically become paralectotypes (art. 73.2.2 of ICZN).
by a single specimen from Djebel Ouach, with lateral tubercles. The specimen figured by Heinz (1886) corresponds to this latter tuberculate form that Sayn (1891) and Fallot and Termier (1923) considered to be the same as H. geronimae (Hermite), although this view was not shared by other authors like Kilian (1910) and Joleaud (1912). In any case, the great majority of the specimens labelled as Am. metamorphicus in the original material, including the lectotype here chosen, is very different from this tuberculate form and closely resemble the specimen figured by Heinz (1886) as Am. diversecostatus. As a consequence, most of the specimens referred to H. diversecostatus by the subsequent authors, who in the main have followed the Heinz interpretation, correspond in fact to H. metamorphicus. 3.12. Distribution This species is a common component of the ammonite assemblages of the upper part of the K. compressissima Zone and the lower part of the Moutoniceras moutonianum Zone (Lower Barremian). It has been found in Algeria, Spain, France, Austria, Rumania and Crimea.
3.9. Dimensions Lectotype. GMH K8086a. D: 17; Wh: 8.2; Wb: 7.6; Wb/Wh: 0.92; U: 3 Paralectotype 1. GMH K8087a. D: 13; Wh: 6.5; Wb: 6.2; Wb/Wh: 0.95; U: 2 Paralectotype 2. GMH K8127a. D: 13.5; Wh: 6; Wb: 6.5; Wb/Wh: 1.08; U: 2 Paralectotype 3. GMH K9163a. D: 12.5; Wh: 6; Wb: 6.5; Wb/Wh: 1.08; U: 2 3.10. Descriptions 3.10.1. Lectotype “GMH K8086a” Small, involute form with subcircular, slightly compressed whorl section with the maximum whorl breadth at midflank. The ornamentation consists of very fine, crowded and flexuous primary ribs arising from the umbilical seam. Primary ribs bi- or trifurcate at midflank and also intercalated ribs can be observed. Some of the secondary ribs join by twos at a small ventrolateral tubercle. The venter is broad, flattened and smooth at this stage. 3.10.2. Figured paralectotypes GMH K8087a. Similar to the lectotype, but with coarser ornamentation at the same diameter. GMH K8127a. Similar to GMH K8087a, but corresponding to a more advanced ontogenetic stage. Tubercles are very well developed and ribs begin to cross the venter. GMH K9163a. Similar to the lectotype, with a slightly more circular whorl section. We also figure a specimen (GMH K8127b, Plate 1, Fig. 1I and J), that we attribute to this species with certain reservations. It is similar to the strongly ornamented morphotypes but, in addition, it shows narrow, periodical constrictions adapically flanked by very strong ribs. This specimen is figured as H. cf. metamorphicus. 3.11. Discussion As we can be seen above, the original diagnosis is quite vague and could be applied to many different holcodiscids. Moreover, Coquand (1880) also included in this species a variety, represented
Family Pulchelliidae Douvillé, 1890 Genus Heinzia Sayn, 1891 Type species: Heinzia sayni Hyatt, 1903 Heinzia heinzi (Coquand, 1880) Plate 1, Fig. 1N and O * 1880 Ammonites Heinzi Coquand, p. 18 non 1886 Ammonites Heinzi Coquand e Heinz, pl. 1 non 1891 Pulchellia Heinzi Coquand in Heinz e Sayn, p. 159; pl. 2, fig. 5aec ? 1891 Pulchellia subcaicedi nov. sp. e Sayn, p. 163, pl. 2, fig. 6 non 1966 Pulchellia heinzi (Coquand) e Breskovski, p. 108; pl. 5, fig. 5 non 1967 Pulchellia heinzi (Coquand) e Dimitrova, p. 166; pl. 80, fig. 6 (¼Breskovski, 1966, pl. 5, fig. 5) 1980a Pulchellia (Heinzia) heinzi (Coq.) e Vermeulen, pl. 4, figs. 11e14 1980b Pulchellia (Heinzia) heinzi (Coq.) e Vermeulen, pl. 6, figs. 11e14 (¼Vermeulen, 1980a, pl. 4, figs. 11e14) non 1995 Coronites (Curiolites) aff. heinzi (Coquand) in Sayn e Vermeulen, pl. 2, fig. 9 non 1997 Curiolites aff. heinzi (Sayn) e Vermeulen, pl. 3, fig. 12 (¼Vermeulen, 1995, pl. 2, fig. 9) 1998a Heinzia gr. sayni Hyatt e Vermeulen, pl. 4, figs. 4e6 (¼Vermeulen, 1980a, pl. 4, figs. 12e14) non 2003 Curiolia heinzi (Sayn) e Vermeulen, p. 158; pl. 41, figs. 5 (¼Vermeulen, 1995, pl. 2, fig. 9), 9e11 2003 Heinzia sp. gr. sayni Hyatt e Vermeulen, pl. 47, fig. 4 (¼Vermeulen, 1980a, pl. 4, fig. 14 3.13. Original diagnosis “Diamètre: 0.017 m. Coquille de petit taille, un peu comprimée, ornée par tours de quatorze côtes simples quelquefois, mais plus souvent bifurquées, partant de l’ombilic, espacées, plates et épaisses, s’arrêtant au milieu du dos, où elles s’infléchissent légèrement en avant en se terminant par un tubercule obtus et tronqué; dos creusé par un sillon profond et étroit, dominé par les tubercules de côtes. Spire formée de tours convexes légèrement anguleux, apparents dans l’ombilic sur la moitié de leur largeur. Bouche subquadrangulaire”.
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3.14. Type material
3.18. Distribution
The only specimen of this species remaining in the Coquand Collection (GMH K8029) is smaller than that originally described by Coquand (1880) and comes from a different locality. Nevertheless, it fits well the original description. We here designate it as lectotype of the species. According to the original label of Coquand, it comes from Djebel Nador. Another specimen, labelled GMH K8097, was a repositored item and had been listed in 1984. It probably was the specimen described by Coquand (1880). Unfortunately all our efforts to find this specimen now have been failed. Vermeulen (2003) designated the specimen of Sayn (1890, pl. 2, fig. 5) as lectotype of C. heinzi Sayn (non Coquand), as he correctly thought that the figured specimen by Sayn did not belong to the same species as those described by Coquand (1880). Therefore, the designation of the lectotype by Vermeulen for C. heinzi Sayn (non Coquand) is not relevant for this paper.
All the specimens that can be undoubtedly attributed to this species (see synonymy list) come from the Barremian of Algeria. Its exact stratigraphic position is not known, but its ornamentation is similar to that of other pulchelliids from the M. moutonianum Zone. Undetermined Family
3.15. Dimensions Lectotype GMH K8029. D: 11; Wh: 5; Wb: 4; Wb/Wh:0.85; U: 2
3.16. Description Tiny, pyritic, rather involute and compressed nucleus of 11 mm diameter. The whorl section is subrectangular, with slightly convex sides and the maximum breadth at midflank. The ornamentation consists of broad, flat ribs that arise, mostly in pairs, from an umbilical swell and develop a strong, truncated tubercle on the outer lateral region. The ventral region is smooth, narrow and concave.
3.17. Discussion Heinz (1886) figured as belonging to this species a specimen that did not correspond to the Coquand description. In the Heinz specimen most of the ribs are simple and rigid, and the ventrolateral tubercles are nodular. Sayn (1891), even recognizing these differences, followed the Heinz interpretation. Furthermore, he described a new species, P. subcaicedi, which showed strong agreement with the Coquand diagnosis of “Ammonites heinzi”. On the other hand, Vermeulen (1980a, b) attributed two specimens to Pulchellia (Heinzia) heinzi from Djebel Ouach that closely match the original description and the specimen held in the Coquand Collection. More recently, however, Vermeulen (1998a, 2003) and Vermeulen and Klein (2006) have related these specimens to H. sayni Hyatt, despite the fact this latter species has grooved ventrolateral tubercles. Meanwhile, Vermeulen (1995) defined a new genus, Curiolites, for the species “Ammonites heinzi” Coquand (1880). Shortly afterwards, however, the same author (Vermeulen, 1997, p. 103) stated that the type species of Curiolites was C. heinzi (Sayn) non Coquand (1880), since he considered that the Coquand species (i. e. heinzi) could be different and should be abandoned. This change is, evidently, against the ICZN rules, as is also the subsequent replacement (Vermeulen, 1999), without any apparent reason, of the name Curiolites by Curiolia. In our opinion, the Coquand species is perfectly valid and finds its place in the genus Heinzia, near other species, like Heinzia caicedi (Karsten, 1858) and Heinzia hispanica Hyatt, 1903 with flat, rhomboidal tubercles.
Undetermined Genus Ammonites mazuca Coquand, 1880 * 1880 Ammonites Mazuca Coquand, p. 368
3.19. Original diagnosis “Diamètre: 0.015 m. Coquille discoïdale, très-comprimée, tranchante et entière à son pourtour, marquée en travers de très légères côtes, plates, flexueuses, distantes, presque sans relief, qui partent du pourtour de l’ombilic, infléchies d’avant en arrière, disparaissant près de la carène. Spire embrassante. Ombilic à peine indiqué. Tours très comprimés. Bouche en fer de lance, très-comprimée, anguleuse en avant, fortement échancrée en arrière”.
3.20. Type Our efforts to find the type material of this species in the collections of the Hungarian Geological Institute have been in vain. Prof. W. J. Kennedy (Oxford University) kindly provided us a list from 1984 of the Cretaceous ammonite specimens of the Coquand Collection. In 1984 he visited the Hungarian Geological Institute for the purpose of research on the Coquand Collection, and the former }-Czabalay gave him the document containing the curator, L. Benko specimens existing at that moment. In the copy, a “Saynella mazuca Coquand” from Djebel Ouach, with the repository number K8079 is clearly indicated. Since then, the museum collections were rearranged, and the Coquand material was temporarily moved to countryside store-houses and then back again. Many specimens were lost in the meantime, and this seems to be also the case of the “Ammonites mazuca”. Vermeulen (1996, 1998b) mentioned that the type material of this species was conserved in the Coquand Collection at Budapest, but without giving any further information. More recently, however, Vermeulen (2003) has designated as lectotype a specimen from his own collection, which contravenes the Article 74 of the Code.
3.21. Discussion After its creation, this species name had never been used until Vermeulen (1996) considered it to be a senior synonym of a wellknown pulchelliid from the lowermost Barremian, Psilotissotia chalmasi (Nicklès, 1890). Since then, Psilotissotia mazuca has been repeatedly used by our colleague to characterize a unit of his successive biostratigraphic schemes for the Barremian of SE France (Vermeulen, 1996, 1998a, 2003, 2005). Nevertheless, we think that there are strong reasons to regard this species as a nomen dubium. Firstly, as stated above, its type material seems to be definitively lost. Secondly, the original description is quite imprecise and could correspond not only to a Psilotissotia, but also to different species of Arnaudiella, Torcapella, Barremites or the Aptian Pseudosaynella or Aconeceras. And lastly, no other of the ammonite species reported from Djebel Ouach (Coquand, 1880; Sayn, 1890, 1891, 1896; Joleaud, 1901, 1912;
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Busnardo, 1957; Vermeulen and Lahondère, 2008) can be exclusively attributed to the lowermost Barremian, and most of them are characteristic of the middle and upper parts of the stage (K. compressissima Zone or higher stratigraphic levels). Under these circumstances, the synonymy between “Ammonites mazuca” and P. chalmasi cannot be verified, so the use of the nominal species proposed by Coquand should be avoided.
4. Summary and conclusions Coquand (1880) established several new species of Barremian ammonites with only short descriptions and no figures, which rendered their identification difficult and controversial. The analysis, for the first time after its original description, of the Coquand’s type material, held in the Hungarian Geological Institute, has enabled us to designate the lectotypes and revise the current conceptions of some of these species. H. diversecostatus, H. metamorphicus and H. heinzi have been misidentified by most of the authors, who have adopted the interpretation by Heinz (1886) of these species. H. diversecostatus (Coquand) is here considered to be a senior synonym of H. nicklesi (Karakasch), whereas most of the specimens attributed by the authors to H. diversecostatus correspond, in fact, to true H metamorphicus (Coquand). In turn, H. heinzi (Coquand) has to be regarded as a different species from the forms figured by Heinz (1886) and Sayn (1891) under this name. Here we also designate lectotypes for H. diversecostatus, H. metamorphicus and Hz. heinzi. The type material of “Ammonites mazuca” has not been found. This and the uncertainty on its stratigraphic position lead us to consider this species name as a nomen dubium, whose application should be fully avoided.
Acknowledgements This paper is a contribution to Project HH 2008-0014 (Hungarian-Spanish Cooperation Programme). The research of M. Company has been also co-financed by Project CGL 2008-00533 (Spanish Ministry of Science and Technology) and Research Group RNM-178 (Junta de Andalucía). The scientific research of O. Szives on the Coquand Collection was supported by the Hungarian OTKA K62063 research project and the TéT ES 28/2008 HungarianSpanish Cooperation Programme. The authors are also grateful to Prof. W. J. Kennedy (Oxford University Museum) for providing them his notes on the Coquand Collection.
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