Origin of the Tethyan Hemihoplitidae Tested with Cladistics (Ancyloceratina, Ammonoidea, Early Cretaceous): an Immigration Event? Didier Bert, Stéphane Bersac

Home , Ancyloceras, Ancyloceratina, Ancyloceratoidea, Crioceras (cephalopod), Crioceratites, Emericiceras

Origin of the Tethyan Hemihoplitidae tested with cladistics (Ancyloceratina, Ammonoidea, Early Cretaceous): an immigration event? Didier Bert, Stéphane Bersac

To cite this version:

Didier Bert, Stéphane Bersac. Origin of the Tethyan Hemihoplitidae tested with cladistics (Ancylo- ceratina, Ammonoidea, Early Cretaceous): an immigration event?. Carnets de Geologie, Carnets de Geologie, 2014, 14 (13), pp.255-272. insu-01071656

HAL Id: insu-01071656 https://hal-insu.archives-ouvertes.fr/insu-01071656 Submitted on 17 Oct 2014

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

Origin of the Tethyan Hemihoplitidae tested with cladistics (Ancyloceratina, Ammonoidea, Early Cretaceous): an immigration event?

Didier BERT 1, 2

Stéphane BERSAC 2

Abstract: The Late Barremian Hemihoplitidae (Ancyloceratina, Ammonoidea) are widely known in the northern Tethyan Margin and the Essaouira-Agadir Basin (Morocco). Their rapid evolution and diversification make them one of the key groups for that period, but their origin remains poorly known and several competing hypotheses have been published. These hypotheses are tested here with cladistic analysis in order to reject those receiving the least support and discuss those well supported. The analysis discards the Crioceratitidae, Emericiceratidae (Emericiceras and Honnoratia) and Toxancyloceras as stem-groups of the Hemihoplitidae (Gassendiceras). The Toxancyloceras appear instead to be a sister-taxon of the Moutoniceras, so we propose the latter to be classified with the Ancyloceratidae rather than with the Heteroceratidae. The best supported hypothesis assumes that the Hemihoplitidae first appeared suddenly in the Essaouira-Agadir Basin at the end of the Early Barremian from small populations of Boreal Paracrioceras. These latter could have migrated southward episodically before invading the northern Tethyan margin at the beginning of the Late Barremian. As a consequence, the Paracrioceratidae fam. nov. is proposed to include the Boreal groups Fissicostaticeras / Paracrioceras / Parancyloceras, and Gassendiceras essaouirae sp. nov. is proposed as a new name for the Moroccan endemic "Barrancyloceras" maghrebiense sensu COMPANY et al., 2008, non IMMEL, 1978. Key Words: Ammonites; Hemihoplitidae; Barremian; cladistics; allopatry; founder effect; palaeobio- geography. Citation: BERT D. & BERSAC S. (2014).- Origin of the Tethyan Hemihoplitidae tested with cladistics (Ancyloceratina, Ammonoidea, Early Cretaceous): an immigration event?.- Carnets de Géologie [Note- books on Geology], Brest, vol. 14, n° 13, p. 255-272. Résumé : L'origine des Hemihoplitidae téthysiens testée par la cladistique (Ancyloceratina, Ammonoidea, Crétacé inférieur) : un événement migratoire ?.- Les Hemihoplitidae du Barré- mien supérieur (Ancyloceratina, Ammonoidea) sont connus sur la plus grande partie de la marge Nord Téthysienne et dans le Bassin de Essaouira-Agadir (Maroc). Leur évolution et leur diversification rapide en font un des groupes clés pour cette période, mais leur origine reste peu connue et plusieurs hypo- thèses contradictoires ont été développées dans la littérature. Ces hypothèses sont testées ici par une analyse cladistique afin d'écarter les moins étayées d'entre elles et de discuter les meilleures. L'analyse écarte les Crioceratitidae, les Emericiceratidae (Emericiceras et Honnoratia) et les Toxancyloceras en tant que groupe-souche pour les (Gassendiceras). D'autre part, les Toxancyloceras apparaissent être taxon-frêre des Moutoniceras, aussi est-il proposé de classer ces derniers parmi les Ancyloceratidae plutôt que dans les Heteroceratidae. L'hypothèse la mieux soutenue suggère que les Hemihoplitidae seraient initialement apparus soudainement à la fin du Barrémien inférieur dans le Bassin d'Essaouira- Agadir à partir de petites populations de Paracrioceras boréaux. Ces derniers auraient pu migrer épisodiquement vers le Sud avant d'envahir la marge Nord-Téthysienne au début du Barrémien supérieur. En conclusion, la Famille des Paracrioceratidae fam. nov. est proposée pour individualiser le groupe boréal Fissicostaticeras / Paracrioceras / Parancyloceras, et Gassendiceras essaouirae sp. nov. est proposé comme nom nouveau pour désigner les "Barrancyloceras" maghrebiense sensu COMPANY et al., 2008, non IMMEL, 1978, endémiques du Maroc. Mots-clefs : Ammonites ; Hemihoplitidae ; Barrémien ; cladistique ; allopatrie ; effet fondateur ; pa- léobiogéographie.

1. Introduction represents a separate entity from the Ancyloce- ratidae GILL, 1871. Their rapid evolution and The marine Late Barremian (pro parte) am- diversification make it one of the key groups for monite family Hemihoplitidae SPATH, 1924 that period. The Hemihoplitidae are currently (Ancyloceratina WIEDMANN, 1960), is widely under revision (D.B.) and several contributions known in the northern Tethyan Margin and the have already been published (BERT & DELANOY, Essaouira-Agadir Basin (Morocco), where it

1 UMR CNRS 6118 Géosciences, Université de Rennes 1, campus Beaulieu, bâtiment 15, F-35042 Rennes 1 cedex (France) [email protected] 2 Laboratoire du Groupe de recherche en Paléobiologie et biostratigraphie des Ammonites (G.P.A.), Bois-Mésange, quartier St Joseph, F-04170 La Mure-Argens (France) Published online in final form (pdf) on October 14, 2014 [Editor: Michel MOULLADE; language editor: Robert W. SCOTT; technical editor: Bruno GRANIER]

255 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

2000, 2009; BERT, 2012, 2013; BERT & BERSAC, phylogenetic relationships. The genus 2012, 2013; BERT et al., 2006, 2008, 2009, Gassendiceras spans the Vandenheckei and 2010, 2011, 2013). These works helped to Sartousiana zones. This latter genus is recognise three major developments for this characterized by a particularly recognizable family. These developments are taxonomically morphology with an uncoiled shell and main characterized by three subfamilies (Fig. 1), trituberculate ribs strongly marked alternately from which the Gassendiceratinae BERT et al., with spineless intercalary ribs (smooth). The 2006, represents the stem of the whole group; Pseudoshasticrioceras are derived from the the group is present from the early Late Gassendiceras in the Feraudianus Horizon or a Barremian Vandenheckei Zone to the Giraudi little earlier. They show a tighter coiling, Zone. depletion of spineless ribs and proliferation of The Gassendiceratinae include the genera weakly tuberculate intermediate ribs (ribs are Gassendiceras BERT et al., 2006, Pseudo- less differentiated from each other on the shasticrioceras DELANOY, 1998, and Imerites ventral margin – BERT & DELANOY, 2009). ROUCHADZE, 1933, which have close

Figure 1: Relationships within the Hemihoplitidae (after BERT, 2012).

256 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

The transition to Imerites (base of the they were supposed to be the descendants. This Giraudi Zone) is accomplished by a major evolution would be carried out through a pro- restructuring of the shell (appearance of the cess of shell recoiling. Such an origination helical early stage), which also determines the would be supported by very strong ornamental morphological appearance of dimorphism (BERT convergences between the juvenile whorls of et al., 2009). Taxonomy, intraspecific variability both the Gassendiceras and Toxancyloceras, and ontogenetic development of the genus which are also present partially in Emericiceras Gassendiceras have been revised recently (alternation of trituberculate main ribs and based on abundant material from south-eastern smooth interribs). For IMMEL (1978, Fig. 11, p. France (BERT et al., 2013; BERT, 2013), and its 70) E. emerici could be directly at the origin of phylogenetic reconstruction was carried out the Gassendiceras (= "E. barremense"). using cladistic analysis (BERT & BERSAC, 2013). 3- Honnoratia thiollierei (ASTIER, 1851) as The results were compared to stem-group: for VERMEULEN (2000, p. 130), the palaeoenvironmental external data (eustacy, Hemihoplitidae originated in the Emericicerati- palaeoclimate proxies) and show that Gassen- dae, but from Honnoratia thiollierei instead of diceras has an anagenetic evolution of palin- the Emericiceras emerici group. This genesis type associated with a constant assumption is based probably on the uncoiling selection under environmental control (trans- and the ornamental similarities between gressive sequence). Honnoratia BUSNARDO et al., 2003, and The origin of the Hemihoplitidae remains Gassendiceras with the trituberculate main ribs poorly known despite this revision, and several alternating with smooth interribs. competing hypotheses have been published. 4- "Barrancyloceras" maghrebiense (sensu The purpose of the present paper is to test the- COMPANY et al., 2008, non IMMEL, 1978) as stem- se hypotheses with cladistics and to resolve the group: Crioceratites maghrebiense IMMEL, 1978, origin of the whole Hemihoplitidae clade, of was classified into the genus Barrancyloceras which the Gassendiceras are the oldest repre- VERMEULEN, 2000 [of which the type species B. sentatives barremense (KILIAN, 1895) is a nomen dubium – see BERT et al., 2010] by KLEIN et al. (2007) 2. Hypotheses on the Hemihoplitidae and COMPANY et al. (2008). Following VERMEULEN origin (note 198, p. 225 in KLEIN et al., 2007), we consider however that the type specimen of the 2.1. The literature hypotheses Moroccan taxon maghrebiense s.s. (sensu IMMEL, non COMPANY et al.) is rather an Early Authors proposed previously four hypotheses Barremian Emericiceras unrelated to the about the origin of the Hemihoplitidae: Hemihoplitidae. According to COMPANY et al. 1- Crioceratitidae GILL, 1871, as stem- (2008, p. 19) "Barrancyloceras" maghrebiense group: for WIEDMANN (1962, p. 112 and Fig. 35), (non IMMEL, 1978 – here renamed into "Hemihoplitidae [are] derived from Pseudothur- Gassendiceras essaouirae sp. nov., see point 6) mannia and perhaps from other Crioceratitidae could be an ancestral endemic Moroccan genera with similar suture". Despite the large representative of the Hemihoplitidae (Agadir- stratigraphic gap between the last Criocera- Essaouira Basin). It characterizes a horizon at titidae (Late Hauterivian) and the first Hemiho- the top of the Moutonianum Zone (end of the plitidae (early Late Barremian), WIEDMANN (follo- Early Barremian). wed by IMMEL, 1979, and more recently by KLIN- Note about the name of T. vandenheckei. As GER & KENNEDY, 1992, and MIKHAILOVA & BARA- previously explained by BERT et al. (2013), the BOSHKIN, 2009) classified the forms of the original spelling (ASTIER, 1851) of this species is "barremense" KILIAN, 1895 / alpinum d'ORBIGNY, Vanden-Heckii. According to the I.C.Z.N article 1850 group (here Gassendiceras) into the 32.5.2.3, this name has to be corrected into genus Crioceratites LÉVEILLÉ, 1837. The former vandenheckii. But this species was dedicated to species are supposed to be derived from the the Abbot VANDEN-HECKE (ASTIER, 1851, p. 452), latter by the increasingly tight coiling of the so the name of the species should be vanden- shell. A neighbouring position was supported heckei. The terminal -ii is incorrect and cor- recently by KAKABADZE and HOEDEMAEKER (2004, responds probably to an inadvertent error (lap- p. 82), who considered the Hemihoplitidae sus calami). The name therefore has to be cor- invalid in favour of the Ancyloceratidae. rected to vandenheckei wherever it appears 2- Emericiceras emerici (LÉVEILLÉ, 1837) and (I.C.Z.N article 32.5.1). Toxancyloceras vandenheckei (ASTIER, 1851) as stem-group: according to DELANOY (1992, p. 2.2. A new hypothesis 52), the Gassendiceras (= group of "E. barremense") would be derived from the Emerici- Following SPATH (1924, p. 82-83) it is possi- ceras SARKAR, 1954, of the E. emerici group ble to point out the very strong morphological (Emericiceratidae VERMEULEN, 2004) via the affinities between Tethyan taxa currently classi- Toxancyloceras DELANOY, 2003, of the T. van- fied in the genus Gassendiceras and the Boreal denheckei group (Ancyloceratidae) of which Paracrioceras of the stadtlaenderi (MÜLLER, 1892) / elegans (von KOENEN, 1902) group, be- 257 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13 cause they share many features. Both genera have in particular the trituberculate main ribs

Figure 2: Comparison of Hauterivian-Barremian Boreal and Thethyan zonations after the data of BODIN et al., 2009, and MCARTHUR et al., 2004 (redrawn from Fig. 6 in BODIN et al., 2009, modified), with long-term trends in belemnite oxygen isotope records from the Vocontian Trough (Tethyan Realm, France) and the Speeton area (Boreal Realm, England). The two major negative shifts (i.e., warmer temperatures) of the Early Hauterivian and Early Barremian are highlighted by grey shading bands. The datum of both stratigraphic scales is the base of the Boreal Elegans Zone, and the green area highlights its correspondence with the Tethyan stratigraphic scale (BERT et al., 2008). The blue arrow is the abrupt and temporary temperature decrease at about the time of onset of volcanism on the Ontong Java Plateau described by MCARTHUR et al. (2004); the red arrow marks the onset of Gassendiceras in the Essaouira-Aga- dir Basin (Morocco) with G. essaouirae sp. nov. The stratigraphic positions of Paracrioceras stadtlaenderi (Boreal Realm, see KAKABADZE & HOEDEMAEKER, 2010), G. essaouirae sp. nov. (Essaouira-Agadir Basin, see COMPANY et al., 2008) and G. multicostatum (Tethyan Realm, see BERT et al., 2013) are also shown; the orange arrows propose the migratory hypotheses, first from the Boreal Realm to the Essaouira-Agadir Basin, and secondly from the latter to the Tethyan Realm (see text). (ventrolateral, lateral and umbilical tubercles) Tethyan Vandenheckei and Sartousiana zones – separated by one to two, rarely three or more, see RAWSON, 1983, p. 494, tab. 1; RAWSON, markedly thinner intermediate ribs between 1995, Fig. 2; OGG & HINNOV, 2012). In fact, they every two main ribs (= the Barremense stage). are present from the boundary between the The ventrolateral spines are stronger than the Elegans and the Fissicostatum zones (see KAKA- lateral and umbilical tubercles. Looped ribs bet- BADZE & HOEDEMAEKER, 2010, p. 37 and 40). The ween the lateral and umbilical tubercles may limit between these latter zones is usually also occur in both genera (KAKABADZE & HOEDE- considered to correspond roughly to the Early - MAEKER, 2010; BERT et al., 2013). The interme- Late Barremian boundary (MUTTERLOSE & BÖCKEL, diate ribs bear weak ventrolateral and someti- 1998; MUTTERLOSE & BORNEMANN, 2000). Howe- mes lateral tubercles in some species of Para- ver, according to a more recent study, the early crioceras as they do in Gassendiceras. Elegans Boreal Zone instead corresponds to the The Paracrioceras of the stadtlaenderi / transition between the Compressissima / elegans group are usually known in the Boreal Moutonianum Tethyan zones in the Early Barre- Denckmani Zone (which is placed in the Late mian (BODIN et al., 2009, p. 1260, Fig. 6 – here Barremian and probably correlates with the Fig. 2). Thus, it is reasonable to expect that the 258 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13 first Paracrioceras of the stadtlaenderi / elegans symplesiomophies). We have taken into group arose in the equivalent of the Moutonia- account the morpho-dimentional parameters of num Tethyan Zone. the shell (characters 0-4 and 17-18): the adult Therefore, the genera Paracrioceras and size from small to very large, the general shape Gassendiceras have a stratigraphic distribution of the shell with the coiling modalities (evolute that is highly consistent with ancestor-descen- normally coiled, crioconic, or tripartite), the dant relationship. growth in height whorl, and the width of the whorl section. The ontogenesis concerns 3. Methods characters 5-6 and 16 with the presence / absence of certain ontogenetic stages Cladistic analysis is nowadays one of the recognised classically within the oldest most popular methods used to infer phyloge- Hemihoplitidae (Gassendiceras): the Heberti, netic relationships. Cladistics is however still Barremense and Simplified ornamentation little used in ammonite studies despite that it is stages. The Heberti stage has simple thin ribs now regarded as an available tool for their evo- slightly differentiated and regularly tritubercu- lutionary studies (NEIGE et al., 2007). The late with a sub-octagonal whorl section. The stratigraphic data for ammonites are indeed Barremense stage has strongly trituberculate deemed of such high quality that many authors main ribs, intermediary ribs, and smooth thin consider these sufficient for the reconstruction interribs alternately. The Simplified ornamenta- of phylogenetic patterns using stratophenetic tion stage has mostly simple ribs with reduced methods alone (GUEX, 2006; see ROUGET et al., tubercles (see BERT et al., 2013, for an exten- 2004, and MONNET, 2005, for contradictory sive description of these stages). Special atten- discussions). However, in the case of the tion was finally given to the ornamental Hemihoplitidae the stratophenetic analysis structures (characters 7-15): the shape and failed to find their origin for which there are five position of the spines, tubercles and ribs. When competing hypotheses (see point 2). Therefore, the state of a character is unknown it has been these hypotheses were tested with cladistics coded by [?]. The characters subject to (parsimony analysis) in order to take into variation (i.e., the multistate/polymorphic account solely the relations of similarity among characters) for a given taxon were coded as the taxa, and so to reject the hypotheses such in square brackets with their different receiving the least support and to discuss the states. best supported. 3.2. Outgroup taxon 3.1. Taxa and characters analysed Polarization is necessary in cladistics to We built a taxon-character matrix (Appen- distinguish derived (apomorphic) from primitive dix) with 11 taxa selected from the previous (plesiomorphic) state of a character, so to hypotheses. Gassendiceras multicostatum (SAR- resolve polarization of the cladogram an KAR, 1955) is currently the oldest known Gas- outgroup comparison was performed. Only one sendiceras species from the north Tethyan outgroup taxon has been selected for the Margin, so we chose it to represent the Hemi- analysis because testing the monophyly of all hoplitidae. The genus Pseudothurmannia SPATH, the taxa selected here is not the purpose of the 1923 (Crioceratitidae), to which was added the present work. We only want to test the related genus Sornayites WIEDMANN, 1962, with hypotheses of the origin of one single group robust ornamentation, represents the first (the Hemihoplitidae) and not to resolve hypothesis about the Hemihoplitidae origin: the phylogeny of the whole tested taxa. The one of WIEDMANN. The second hypothesis, the outgroup taxon we chose is the "primitive" DELANOY's, is represented by the genera Eme- Ancyloceratina Crioceratites nolani (KILIAN, riciceras and Toxancyloceras. The genus Honno- 1919), which is a large sized crioconic species. ratia was added to test the third hypothesis of This taxon has also the advantage to be VERMEULEN. Gassendiceras essaouirae sp. nov. undoubtedly older than all the other taxa tested corresponds to "Barrancyloceras" maghrebiense because it appeared in the Early Hauterivian. in sense of COMPANY et al. (non IMMEL – see point 6). Both the Boreal taxa Paracrioceras 3.3. Discussion about the relevancy of stadtlaenderi (MÜLLER, 1892) and the matrix Fissicostaticeras KAKABADZE & HOEDEMAEKER, 2010, represent the last hypothesis (this work). 3.3.1. Discretization of continuous The genus Moutoniceras SARKAR, 1955, was characters added because it is one of the rare Tethyan heteromorphic ammonites with suture line of Some of the selected characters (characters ancyloceratid type at the end of the Early 0, 2, 3 and 4, see Appendix) are objectively Barremian, and therefore it could not be continuous but could not be encoded as such neglected. (see GOLOBOFF et al., 2008; BERT & BERSAC, Nineteen discrete characters were selected 2013) because of the lack of revision of the ma- (see Appendix) after having excluded the unin- jority of the studied groups, which limits the formative characters (autapomorphies and numbers of reliable measurements to extract 259 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13 the phylogenetic information. In the current problematic if it belongs to the ingroup of the state of knowledge, we considered that estima- study (BRYANT, 1997). In the present paper, we tion is less reliable than a discretized value, so chose to treat the supraspecific taxa according we performed discretization using the classical to the exemplar method as described by PREN- method: the difference between the maximum DINI (2001 – one or more of its representative and minimum observed or estimated values species replaced each generic taxon of the ma- was divided by the number of states for the trix). Selecting a representative species usually characters 0, 2 and 3. For the character 4 consists in choosing the most common or the (coiling hiatus), this range was divided by 2, supposed earliest / ancestral species (BININDA- because the "first" character-state corresponds EMONDS et al., 1998; PRENDINI, 2001). However, to a value of 0. in the present study this method is problematic because the relative abundance and the evolu- 3.3.2. Treatment of polymorphic tionary patterns of the treated taxa are not pre- characters cisely known (non-revised taxa most of the The matrix contains eight characters (42%) time, cf. supra), and because their monophyly with a polymorphic state (see Appendix). They has never been tested. In other words, the risk represent 8.3% of the total phylogenetic infor- here is to select an inappropriate species, which mation and 8.2% of the total data (the four un- does not belong to the genus it is supposed to known states represent 1.8% of the total data). represent. For this study, only one species per Multistate characters coded as such are proble- genus was selected to not overload the clado- matic because they introduce "extra" homo- gram: the type species in order to ensure that plasy in providing soft reversals (according to it belongs for sure to its genus. So the seven taxa at the generic level of the matrix were KORNET & TURNER, 1999, p. 366, "a soft reversal takes place when an unfixed evolutionary no- replaced (see point 3.1) by Pseudothurmannia velty disappears from a polymorphic lineage", picteti SARKAR, 1955, Sornayites paronai (SARKAR, 1955), Emericiceras emerici (LÉVEILLÉ, see KORNET & TURNER, 1999, for explanations) and thus, they should be avoided as much as 1837), Toxancyloceras vandenheckei (ASTIER, possible. Therefore, the polymorphic characters 1851), Honnoratia honnoratiana (d'ORBIGNY, were treated here according to the method 1842), Fissicostaticeras fissicostatum (ROEMER, 1841), and Moutoniceras moutonianum proposed by KORNET and TURNER (1999), which gives the most congruent results. When pos- (d'ORBIGNY, 1850) respectively. After replacing sible, the polymorphic characters were coded each generic taxon by its type species, then according to their ancestral state (Inferring An- applying as far as possible the IAS method for cestral State, or IAS method). In case of impos- polymorphisms (or ambiguous method if not sibility to discriminate the ancestral state from possible), only two characters (10.5%) remain the derived state, the polymorphic characters polymorphic for seven taxa: characters 3 and were coded as such ("ambiguous coding" of 12 (Appendix). As a result of using this method, the proportion of polymorphisms versus the KORNET & TURNER, 1999, p. 370). In fact, most of the polymorphism (94.7%) is because we used total phylogenetic information in the matrix seven taxa of generic rank (= 63.6% of the decreases from 8.3% to 3.9%. We chose not to taxa – Appendix): coding the characters of follow the recommendation of PRENDINI (2001) supraspecific taxa consists of adding the diffe- to represent supraspecific (here genera) taxa rent character states of their species, which in- by several species (including the type species) creases the probability of polymorphic cha- in order to take into account their phenotypic diversity and eventually to test their racters (KRON & JUDD, 1990; NIXON & DAVIS, monophyly. In the present study, as pointed 1991; PRENDINI, 2001). The accuracy of the numerous methods available to treat polymorphic above, the "non-type" species may not belong characters of supraspecific taxa in cladistics has to their genus for sure. Therefore, including such species in the cladistic analysis would long been debated (see PRENDINI, 2001, and potentially add useless information, because SIMMONS & GLEISER, 2002, for a discussion). From one method to another, the supraspecific the goal of the present paper is not to test the taxa could be replaced in the matrix by one monophyly of the generic selected taxa. species or specimen (exemplar method), or the 3.4. Analysis method polymorphic characters could be coded sepa-ra- tely according to one state only (the ancestral The analysis was performed with the TNT state or the most frequent state). The separate software (Tree analysing using New Tech- coding of polymorphic characters (e.g., IAS / nology), version 1.1 for Windows (GOLOBOFF et ancestral / compartmentalization, democratic al., 2008) using the "branch-and-bound" me- methods, see KORNET & TURNER, 1999; PRENDINI, thod via the Implicit Enumeration option 2001, for a short description) can bias the (collapsing rule used is maximum length = 0) results of the cladistic analysis because it pre- that allows finding for sure the most parsimo- sents the risk in providing a chimeric taxon (see nious trees. The characters were considered PRENDINI, 2001). Such a chimeric taxon, with all unordered and unweighted in order to avoid the its polymorphic characters "forced" to their an- inference of pre-analytical assumptions. The cestral state (i.e., a hypothetical ancestor), is Consistency Index (KLUGE & FARRIS, 1969), the 260 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

Retention Index (FARRIS, 1989) and the Ad- uninteresting but it has to be considered justed Homoplasy (GOLOBOFF et al., 2008) quan- more carefully. Nevertheless, a BREMER value of tified homoplasy. In case of multiple most 1 can be used to assess and reinforce the parsimonious trees, a strict consensus clado- interest of this node. gram was computed in order to analyse consistency between the trees. Then the eventual ambiguous apomorphies were performed by ACCTRAN optimization (accelerated transforma- tion – FARRIS 1970; SWOFFORD & MADDISSON, 1987, 1992), which favours reversals over convergences. A resampling technique using bootstrap method (standard bootstrap of TNT with 1000 re- plications, using implicit enumeration, collapsing groups below 1%, result given in absolute frequency, see GOLOBOFF et al., 2008) and a decay analysis (BREMER, 1994, Absolute BRE- MER Support in TNT, support calculation with Tree Bisection and Reconnection from existing trees, retained suboptimal trees by 20 steps and null relative fit difference, see GOLOBOFF et al., 2008) were performed with TNT in order to assess the confidence of the nodes and to test the robustness of the tree topology. The bootstrap indicates the stability of the most parsimonious clades under random weighting of characters. So the more this value is high means the more times a given branch occurs in the consensus bootstrap trees, and the more robust this clade is assumed to be. The BREMER support indicates the robustness of a clade by calcula- ting the number of extra steps needed to col- lapse this clade.

4. Results

4.1. The most parsimonious trees The analysis results in 5 most parsimonious trees (Fig. 3) of 40 steps each, which reveal the presence of homoplasies (Consistency Index = 0.675; Retention Index = 0.75; Adjusted Ho- moplasy = 2.95 to 3.05). These five trees have some important topologic similarities since some taxa are systematically paired as sister taxa. This is the case for Toxancyloceras vandenheckei and Moutoniceras moutonianum, Sornayites paronai and Pseudothurmannia picteti, Emericiceras emerici and Honnoratia thiollerei and the two species of Gassendiceras. The other most important similarity between the trees is the clade with Paracrioceras, Fissicostaticeras and the two species of Gassendiceras, which is systematically present in the same structure.

4.2. The strict consensus tree The structure of the strict consensus tree is given in Figure 4. Most of the nodes receive bootstrap value around or greater than 70% and a significant BREMER support value (from 1 Fig. 3: The five most parsimonious trees obtained at to 2 – Fig. 4), which values are overall coherent first run with TNT. between each other. The node with bootstrap value lesser than this threshold (node 17) is not

261 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

The consensus tree shows four major clades 15) and the presence of fibulate ribs are the well separated within the ingroup from a clearly synapomorphic characters supporting this unresolved node (node 13 – Fig. 4), which con- clade. The very large adult size is homoplasic tains no synapomorphy in our study. This result (see Fig. 5). There are five convergent is quite logical due to the spread choice of the characters (homoplasies) between the taxa belonging to different families. The four Honnoratia and the clade of the two species of clades are as follow: the one consisting of the Gassendiceras (characters 4, 9, 11, 14, 17 – Emericiceratidae (Emericiceras emerici and see Appendix). Therefore, the hypothesis of an Honnoratia thiollierei); the one consisting of the origin of the Hemihoplitidae within the Crioceraritidae (Pseudothurmania picteti and Honnoratia is problematic in terms of mor- Sornayites paronai; the pair Moutoniceras and phology (shape of the shell and ornamenta- Toxancyloceras; and the clade consisting of Pa- tion). It is also problematic considering the racrioceras / Fissicostaticeras and Gassendice- stratigraphic gap of nearly two ammonite zones ras. This latter clade has a pectined structure (Compressissima and Moutonianum zones) bet- (each node gives birth to a single taxon and a ween the last Honnoratia (Early Barremian) and clade), with the respective apparition going up- first Gassendiceras (early Late Barremian or ward although the cladogram of Fissicostatice- extremely late Early Barremian according to ras fissicostatum, Paracrioceras stadtlaenderi COMPANY et al., 2008). and the two species of Gassendiceras. This lat- 3- Emericiceras emerici and Toxancyloceras ter clade is the most robust of the tree (Fig. 4). vandenheckei as stem-group (the DELANOY hypothesis): the link DELANOY had evoked between the Emericiceras and Toxancyloceras is not recognised from the cladistic analysis, as the "group" forming these two genera is clearly polyphyletic and thus unacceptable. Emericice- ras emerici is the sister taxon of Honnoratia thiollerei (both Early Barremian Emericicerati- dae) as seen above, and they have relationship with neither the Hemihoplitidae nor the Toxan- cyloceras (Ancyloceratidae). Moreover, this latter genus appears to be sister taxon of Mouto- niceras. The synapomorphic character of the clade supported by Toxancyloceras and Mouto- Fig. 4: Strict consensus tree computed with statistics niceras is the strict ancyloceratic coiling (Fig. (Bootstrap values and BREMER support) (Consistency 5). The coil of some robust Toxancyloceras of Index CI = 0.643; Retention Index RI = 0.712; Ad- the T. vandenheckei group may resemble some justed homoplasy Aj = 3.45). Gassendiceras to a strictly ornamental point of view, especially when they have many 5. Discussion inermous (smooth without any tubercles) interribs. Toxancyloceras and Gassendiceras It is possible to discuss the hypotheses share common characters: their adult size is exposed above (point 2) in light of the similar (character 0); the ribs can be reduced cladogram obtained. on the venter (character 8), are wedge shaped 1- Crioceratitidae as stem-group (the WIED- (character 13), and they often bear robust MANN hypothesis): the adult size, the evolute tubercles (character 15). The flanks are coiling and the shape of the dorsum are the sy- rounded in the same way (character 17). These napomorphic characters supporting the clade of similarities are however convergences Pseudothurmannia / Sornayites (node 14 – Fig. (homoplasies) with no phyletic link, and in 5). The other characters are all homoplasic. It Toxancyloceras the uncoiling is more appears in fact that the hypothesis of the Crio- pronounced with a higher spiral gap (characters ceratitidae as stem of the Hemihoplitidae is the 1 and 4), growth in height whorls is lower result of a typological conception of both the (character 2), and the smooth interribs are still Late Hauterivian Crioceratitidae and the Barre- more numerous (character 10). The shaft of mian Hemihoplitidae based on morphological Toxancyloceras is also more slender, straight convergences (homoplasies) without any phyle- and longer in proportion than in the few tic link between these groups. The hypothesis tripartite species belonging to the genus Gas- of such a link is anyway hampered by the very sendiceras (see BERT et al., 2013). The hook is large stratigraphic gap between the last Pseu- longer, rounded and less tuberculate near the dothurmannia (latest Hauterivian) and the first ventral marginal area in Toxancyloceras. The Hemihoplitidae (Late Barremian). possibility of an origin of the Gassendiceras 2- Honnoratia thiollierei as stem-group (the within the Moutoniceras is utterly inconceivable VERMEULEN hypothesis): Honnoratia thiollierei because of their strongly divergent morphology, appears to be a sister taxon of Emericiceras and such a hypothesis would not be supported emerici and they are both grouped into the by the present cladistic analysis anyway. Emericiceratidae. The coiling of the shell (node 262 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

4- Gassendiceras essaouirae sp. nov. (= Palaeogeographically, the Essaouira-Agadir "Barrancyloceras" maghrebiense sensu COMPANY Basin is of intermediate position on the only et al., 2008, non IMMEL, 1978) as stem-group possible migration route (Fig. 6), while any (the COMPANY et al. hypothesis): examination of other communication path between the Boreal casts of the Moroccan specimens figured by and Tethyan realms was closed again in the COMPANY et al. (2008, Pl. 8, figs. L, M) allows Barremian (Neocomian Polish Furrow), or not the consideration of their classification into the yet opened (Russian Platform, Pyrenean genus Gassendiceras close to G. multicostatum. Furrow, Paris Basin, Polish Through – DERCOURT Their uniqueness merits a new species name et al., 2000). This Basin has a distinctly (see below the taxonomic implications), thus G. Mediterranean character (COMPANY et al., 2008), essaouirae sp. nov. would now represent the and the sudden onset of the Hemihoplitidae at oldest known Gassendiceras. G. essaouirae sp. the end of the Early Barremian in Morocco is nov. is sister taxon of G. multicostatum on the thus strongly consistent with the migratory cladogram as the most robust clade (bootstrap hypothesis of this group towards the Tethys. of 86% consolidated with the significant BREMER The beginning of the transgressive system tract support value of 2 – Fig. 4), and this is strongly (HARDENBOL et al., 1998) at the end of the Early in favour of the COMPANY et al. hypothesis. The Barremian / beginning of the Late Barremian synapomorphy that supports this clade is the s.l. also provides an additional argument with scarcity of the intercalatory ribs. The presence the greater ease of communication induced. of the ontogenetic Heberti stage might be a synapomorphy of this clade as well as a synapomorphy of the Paracrioceras-Gassendiceras clade, due to the unknown state of the character 5 (Fig. 5). The origin of the Tethyan Gassendice- ras within the Moroccan species is also stratigraphically consistent because G. essaouirae sp. nov. is from the latest Early Barremian and G. multicostatum (the oldest-known Tethyan Hemihoplitidae) is from the early Late Barre- mian (see Fig. 2). 5- Paracrioceras as stem-group (hypothesis of the present work): in the cladogram, Para- crioceras is the sister taxon of the Gassen- Fig. 5: Strict consensus tree with character state changes labelled (see Appendix for the list of diceras clade. This close relationship between characters). Paracrioceras and Gassendiceras, and both the correspondence between their appearance on On the other hand, a communication bet- the cladogram (starting from the root) and their ween the Boreal Realm and the Atlantic for the stratigraphical distribution (see above point 2.2 "Mittelbarrême" was argued by IMMEL and MUT- and Fig. 2) are consistent with this hypothesis TERLOSE (1978, p. 265). These latter authors (see BERT & BERSAC, 2013, for a discussion also suggested multiple Boreal influences about cladogram and phylogenetic tree). They during the Moroccan Cretaceous [based on are supported by some synapomorphies (as for ROCH's work (1930) who reported several the Fissicostaticeras – node 16 – Fig. 5), espe- Barremian Boreal ammonite taxa in the Agadir cially concerning the ontogeny (the presence of area], although COMPANY et al. (2008) did not the Barremense and the Simplified ornamen- confirm it for the Barremian. tation stages, characters 6 and 16). Therefore, Note that an East communication path (from the link between these taxa is consolidated, and the Boreal Ocean North of the Scandinavian the origin of the Hemihoplitidae is compatible Shield and through all the Russian Platform) with the migration event hypothesis developed has been proposed to explain the presence of above (point 2.2). Paracrioceras denckmani in the early Late The cladistic analysis clearly dismisses the Barremian of Crimea (KAKABADZE, 1981, Pl. 2, first three hypotheses. The two remaining fig. 1, reconsidered as P. cf. elegans in hypotheses are in contrast consolidated. There KAKABADZE, 1983, p. 506; see also DELANOY & is no incompatibility between them and they FÉRAUD, 1995, Fig. 4 and p. 211). However this should be regarded as complementary. It is determination is challenged here because indeed quite possible that the Hemihoplitidae KAKABADZE's specimen instead belongs to appeared in the Essaouira-Agadir Basin at the Hemihoplites feraudianus (robust morphology), end of the Early Barremian (COMPANY et al., which is unknown in the Boreal Realm. So, this 2008), before invading the northern Tethyan theory is not supported by data until the margin at the beginning of the Late Barremian. dissemination of Spinocrioceras polyspinosum They could be derived from small populations of KEMPER, 1973, in the Late Barremian (spread Boreal Paracrioceras, which might have out from the late Sartousiana Zone in western migrated southward episodically. Tethys, to the early Giraudi Zone in Crimea, and to the early Bidentatum Zone in the Boreal area – see DELANOY & FÉRAUD, 1995). 263 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

Palaeoclimatically, at least three positive 6. Taxonomic implications δ18O excursions are between the end of the Early Barremian and the earliest Late 6.1. Classification of the genus Barremian of the Angles section (the Barremian Moutoniceras stratotype in the Tethyan Realm) in a "greenhouse" context (WISSLER et al., 2002; When introduced, the genus Moutoniceras BODIN et al., 2005). Such peaks are usually was classified in the Heteroceratidae SPATH, regarded as significant to a temporary climatic 1922 (SARKAR, 1955, p. 24). More recently this destabilization (cooling). But here most attribution was challenged by some authors. probably, according to the authors (MCARTHUR et Some (e.g., AUTRAN et al., 1986, p. 1060; DELA- al., 2004, p. 265-266), these peaks of high NOY et al., 1991; COMPANY et al., 2008) proposed frequency oscillations are probably due to local a classification in the Ancyloceratidae, while diagenetic overprint. But in their paper others (e.g., KAKABADZE & THIEULOY, 1991) pro- 18 MCARTHUR et al. (2004, p. 269, based on δ O posed a classification in the Crioceratitinae. and Mg proxy) demonstrated that through the WRIGHT et al. (1996, p. 216) more or less Barremian in the Boreal Realm (Eastern accepted Moutoniceras with doubt in the Ancy- England), temperature increased to a peak of loceratidae, but they also suggested that it 20°C in the Elegans Zone then, in the same could be a Crioceratitinae. Finally, VERMEULEN zone, precipitately and temporarily decreased (1997) proposed to reinstate the genus in the to around 14°C at about the time of onset of Heteroceratidae because of strong morphologi- volcanism on the Ontong-Java Plateau, before cal convergences. In fact until 2007 and the returned to around 16°C. As recently stated by work of KLEIN et al. (2007) there was no robust BODIN et al. (2009, p. 1260, Fig. 6 – see Fig. 2 consensus about the classification of the genus and above point 2.2), the early Elegans Boreal Moutoniceras. In the latter work, no less than Zone corresponds to the transition of the nine co-authors, specialists of the Barremian Compressissima / Moutonianum Tethyan zones (including DELANOY, KAKABADZE and VERMEULEN, in the Early Barremian, and thus the Late but excepting COMPANY according to the footnote Moutonianum Zone (where the Gassendiceras No. 130, p. 174), finally considered the genus appear in the Essaouira-Agadir Basin) could be to be a Heteroceratidae. Nevertheless, the pre- contemporary with this temporary cooling. sent cladistic analysis shows Moutoniceras as a Therefore, it might be tempting to hypothe- sister taxon of Toxancyloceras (strict Ancyloce- size that migration of small populations of ratidae). A link between both genera is con- Boreal Paracrioceras could occasionally be esta- sistent, first stratigraphically and secondly mor- blished under favourable conditions during their phologically: in the Vocontian Basin (South-East expansion into the lower latitudes (REBOULET et of France) the genus Moutoniceras disappeared al., 1992; REBOULET & ATROPS, 1995; REBOULET, with the tuberculate M. eigenheeri (VERMEULEN, 2007). Such small populations were able to 2003) at the very end of the Early Barremian, establish themselves locally in the Essaouira- just below the First Apparition Dating (FAD) of Agadir Basin (Atlantic Realm). The exploitation the genus Toxancyloceras (beginning of the of new ecological niches (for example those left Late Barremian). It seems highly probable that vacant by the disappearance of the Emerici- the genus Toxancyloceras originated from Mou- ceratidae a long time before in the Pulchella toniceras, mainly by the acquisition of the lower Zone, and maybe used later by the tubercles, and the generalisation of the median Moutoniceras) would then have allowed local tubercles, on main ribs. The direct link between adaptation leading to the evolution towards the Toxancyloceras and Moutoniceras involves ran- Gassendiceras morphology (allopatry and king the genus Moutoniceras among the Ancy- speciation by founder effect in sense of MAYR, loceratidae rather than in the Heteroceratidae. 1974). Then, the newly emerging The Heteroceratidae are not beyond the scope Hemihoplitidae invaded and diversified through of the present work, and thus they have not the northern Tethyan margin from this been added in the cladistic analysis, however intermediate geographic position at the we can note that the apparition of the primitive beginning of the Late Barremian (Fig. 6). Heteroceratidae was much more discreet and belated. Exceptionally rare fragments of pos- Note also that in the Boreal Realm, during sible Heteroceratidae were reported from the the Denkmanni Zone (Late Barremian), the lower part of the Sartousiana Zone (VERMEULEN, Paracrioceras continue to evolve in northern 1995), but the first true representatives of the Europe towards forms close morphologically to genus Heteroceras d'ORBIGNY, 1850, are only the southern Gassendiceras [e.g., P. tuba (von known with certainty at the end of the Ferau- KOENEN, 1902)], although they lived in different dianus Subzone (Autrani Horizon – see DELANOY, environments. This might suggest the existence 1994; BERT et al., 2008) with Heteroceras coul- of a common gene pool rather than a simple leti DELANOY, 1994. This stratigraphic position transient morphological convergence. makes a gap of more than one and half ammonite zones between the last Moutoniceras and the first known Heteroceras s.s. (i.e., with a known helical part). 264 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

Fig. 6: Palaeogeographic map of the Barremian (reconstructed from BARRON et al., 1981, and DERCOURT et al., 2000) with extension of the Tethyan Gassendiceras and the Boreal Paracrioceratidae fam. nov. 6.2. The Moroccan Gassendiceras Derivation of the name. Because of its origin (from Essaouira). According to the present analysis, it is necessary to recognise under a new name the Holotype. The specimen No. X.OG.R.23 from Tafadna (Morocco), housed in the pa- "Barrancyloceras" maghrebiense sensu COMPANY laeontological collections of the University of et al., 2008, non IMMEL, 1978, which ranks near the Tethyan Gassendiceras. We propose the na- Granada (Spain). Specimen figured by COMPANY me Gassendiceras essaouirae sp. nov. et al. (2008, Fig. 8M). Paratype. The specimen No. X.MS.R.9 from Family Hemihoplitidae SPATH, 1924 Imsouane (Morocco), housed in the palaeonto- Subfamily Gassendiceratinae BERT et logical collections of the University of Granada. al., 2006 Specimen figured by COMPANY et al. (2008, Fig. 8L). Genus Gassendiceras BERT et al., 2006 Type locality. The Tafadna section (co- Gassendiceras essaouirae sp. nov. ordinates 31º05'46"N 9º48'22"W), 16 km NW of Tamanar; outcrop on the right bank of the Synonymy Oued Igouzouln, some 2 km before it joins the non 1978. Crioceratites (C.) maghrebiensis nov. Atlantic Ocean on Tafadna beach, Essaouira sp.: IMMEL, p. 59, tab. 10b, Pl. 8, fig. 1. Basin, Morocco (COMPANY et al., 2008). v 2008. "Barrancyloceras" maghrebiense (IMMEL, Bed type. The bed of the specimen chosen 1978): COMPANY et al., Fig. 8L-M. here as holotype was not indicated by COMPANY

265 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13 et al. (2008), but the species is present in beds known only in Morocco, and only at the extreme 57-58 of the Imsouane section where the para- end of the Early Barremian (see COMPANY et al., type specimen comes from (COMPANY et al., 2008). 2008, Fig. 3). Geographic distribution. Gassendiceras 6.3. The case of Fissicostaticeras / essaouirae sp. nov. is actually only known in Paracrioceras the Essaouira-Agadir Basin (Morocco), Atlantic If we consider only the Boreal Realm, the Realm. genera Fissicostaticeras and Paracrioceras (and Stratigraphic distribution. Uppermost also Parancyloceras SPATH, 1924, not added in Lower Barremian strata dated from the end of the present cladistic analysis – see KAKABADZE & the Moutonianum Zone. Following COMPANY et HOEDEMAEKER, 2010) are a monophyletic clade, al. (2008) this species characterizes the which requires them to be placed into a separa- Moroccan "Maghrebiense Horizon", here te family. However, the present cladistic ana- renamed as the Essaouirae Horizon. lysis resulted in the recognition of a possible Diagnosis. Only the inner whorls are known phyletic link between the Boreal Fissi- up to D = 80 mm (the first whorls are un- costaticeras / Paracrioceras and the Atlantic / known). Massive shell with tight crioconic coi- Tethyan Gassendiceras (Figs. 4 - 5), which ma- ling; whorl section very broad and rounded of kes de facto the former a paraphyletic clade. As sub-octagonal shape at ornamentation. Only explained by BERT and BERSAC (2013) for the one ornamental stage, very close to the Heberti genus Gassendiceras itself, the rise of a new stage, is known: ribs slightly differentiated, group (here Gassendiceras) would not affect trituberculate, radial or slightly retroverted, and the older group (here the phylum sometimes wedge shaped. Tubercles small, Fissicostaticeras / Paracrioceras). This reflects conical and well defined. They are located at the reality of coexistence (even in different the lower and upper thirds of the flanks, and at geographic areas) of different lineages the peri-ventral border. (HÖRANDL, 2007), because of evolutionary processes where descendants exist without Note. The authors of the present work could concomitant extinction of the parental group. observe the presence of the Barremense and Such processes rendering the parental group Simplified ornamental stages (and of the He- paraphyletic (any group of descendants berti stage in the innermost whorls of the shell) automatically cancels out the monophyly of the on complete and unrestored Moroccan speci- ancestral group; see HÖRANDL, 2006). As mens of G. essaouirae sp. nov. and thus coded HÖRANDL and STUESSY (2010) recognised, the matrix of the cladistic analysis accordingly cladogenesis is the main source of paraphyly (see Appendix). It is to note that these speci- and paraphyly is a normal stage in the evo- mens cannot be figured for ethical reasons lutionary process whereby a new species arises. because they were extracted for commercial use and because their trace has been lost since Considering that paraphyletic groups are (pictures of one of them can be sent on acceptable as taxa in evolutionary classifica- demand). tions, the proposition of allopatric origin of the Tethyan Hemihoplitidae from the Boreal Para- Differential diagnosis. The adult develop- crioceras via the Essaouira-Agadir Basin ments of Gassendiceras essaouirae sp. nov. are requires separating taxonomically the Boreal not known on the basis of the type material lineage Fissicostaticeras / Paracrioceras / (COMPANY et al., 2008), but this species is uni- Parancyloceras from the Emericiceratidae in que and different from all the other Gassen- which they are currently classified (cf. KLEIN et diceras described. Even if the innermost whorls al., 2007). Therefore, we propose to introduce a are unknown, the Heberti stage looks to have a new family: the Paracrioceratidae fam. nov. significantly longer duration, at least until D = 80 mm. This falls perfectly within the evolu- Superfamily Ancyloceratoidea GILL, tionary trends of the Tethyan Gassendiceras 1871 defined by BERT and BERSAC (2013). The shell has also a stronger general appearance with Family Paracrioceratidae fam. nov. larger ribs but smaller tubercles. Type genus. Paracrioceras SPATH, 1924. "C. (C.)" maghrebiensis IMMEL, 1978, differs Generic content. This family includes the from G. essaouirae sp. nov. by the presence of Boreal genera Paracrioceras, Fissicostaticeras many smooth interribs between the main ribs. and Parancyloceras in order to recognise their The former bears only small tubercles, which phylogenetic relationships. A Boreal phyletic are better defined in the latter. In addition, lineage Fissicostaticeras -> Paracrioceras -> IMMEL's species shows the ribs projected forward Parancyloceras may be considered taking into on the uppermost part of the flanks, while in G. account the similarity between these genera, as essaouirae sp. nov. the ribs are radial (stronger well as stratigraphic and paleogeographic argu- and more spaced), even on the ventral area. ments (KAKABADZE & HOEDEMAEKER, 2010). Stratigraphically and geographically G. essa- Phyletic position. The Paracrioceratidae ouirae sp. nov. is very unique because it is yet fam. nov. appear from crioconic forms classified 266 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13 in the genus Emericiceras [sic] by KAKABADZE the beginning of the Late Barremian. They may and HOEDEMAEKER (2010) but probably closer to have been derived from small populations of the Crioceratitidae sensu lato. The Paracriocera- Boreal Paracrioceras, which could have tidae fam. nov. seem to be at the origin of the migrated southward episodically. Such a Tethyan Hemihoplitidae (Gassendiceratinae) by migration of the Paracrioceras would have been allopatry and founder effect. enabled by the establishment of favourable Remarks. It should be noted that KAKABADZE climatic conditions during their expansion and HOEDEMAEKER (2010) used the genus Acrio- towards the lower latitudes (an abrupt and ceras HYATT, 1900, in a different way than temporary temperature drop in the middle part French authors. Some specimens they figured of the Elegans Zone). This is supported [e.g., "Acrioceras" of the nodulosum (von KOE- paleogeographically because the Essaouira- NEN, 1902) group] could possibly correspond to Agadir Basin is of intermediate position on the microconchs of the contemporary Paracrioce- only possible migration route, the Atlantic one ratidae fam. nov. (same ornamental evolution (Fig. 6), in times of transgressive shorelines over time). This hypothesis was suggested by that probably induced a greater ease of RAWSON (1975, p. 282) about the small tripar- communication. The exploitation of new tite genus Hoplocrioceras SPATH, 1924, which ecological niches (for example those left vacant could correspond possibly to the microconch of by the disappearance of the Emericiceratidae) Fissicostaticeras (= Paracrioceras for RAWSON). would then have allowed local adaptation These small tripartite forms would then have to leading to the evolution towards the Gassen- be included in the Paracrioceratidae fam. nov. if diceras morphology (Hemihoplitidae), which this dimorphism was demonstrated. Such a di- invaded and diversified through the northern morphism had been comparatively successfully Tethyan margin. Therefore, the origin of the advanced for the Crioceratitidae by DELANOY et Hemihoplitidae looks to be a case of allopatry al. (1995), and ROPOLO and GONNET (1995), and and speciation by founder effect. for some Hemihoplitidae (DELANOY et al., 1995; Migration from the Boreal Ocean towards the BERT et al., 2009; BERT, 2012, 2013). Essaouira-Agadir Basin is supported morphologi-cally by cladistics and by other arguments, 7. Conclusions but of course, it remains a hypothesis that needs to be tested more thoroughly, especially The marine Late Barremian ammonite family because there is no evidence currently of other Hemihoplitidae (Ancyloceratoidea) is diversified faunistic Boreal influence in Morocco during the in most of the northern Tethyan Margin and the early Late Barremian. Maybe in the future a Essaouira-Agadir Basin (Morocco). However, better competitor than the Paracrioceratidae their origin remained hardly known because nov. fam. could be found to explain the origin several competing hypotheses have been deve- of the Hemihoplitidae? Meanwhile, it remains loped in literature until recently. Most of the the best candidate. literature hypotheses were rejected by cladistics (Crioceratitidae, Emericiceratidae and Toxan- Acknowledgements cyloceras as stem-group) and our analysis shows that they were in fact the result of typo- The authors warmly thanks Miguel COMPANY logical conceptions. Gassendiceras essaouirae and Gérard DELANOY for sending us the casts of sp. nov. appears now to be the best candidate G. essaouirae sp. nov. Special thanks are due as an ancestral endemic Moroccan representa- to Steve FRIEDRICH who kindly improved our ori- tive of the Hemihoplitidae at the end of the ginal English text. We also acknowledge the Moutonianum Zone (end of the Early Barre- three reviewers, Miguel COMPANY, Jérémie BAR- mian). Cladistics also supports an origination of DIN and an anonymous reviewer, for their con- the Hemihoplitidae (incl. G. essaouirae sp. structive remarks, which helped us to improve nov.) from the Boreal Paracrioceratidae fam. the manuscript. nov. (migration hypothesis). Paracrioceras of the stadtlaenderi group have indeed very strong Bibliographic references morphological affinities with the Tethyan Gas- ASTIER J.-E. (1851).- Catalogue descriptif des sendiceras. Stratigraphically, the former is Ancyloceras appartenant à l'étage Néoco- known beginning at the boundary of the Fissi- mien d'Escragnolles et des Basses-Alpes.- costatus / Elegans Boreal zones (late Early Bar- Annales des Sciences Physiques et Natu- remian). Because it is recognised that the Ele- relles, d'Agriculture et d'Industrie, Lyon, gans Zone corresponds roughly to the Mouto- (2ème Série), t. III, p. 435-456. Online at nianum Tethyan Zone, this distribution is http://gallica.bnf.fr/ark:/12148/bpt6k134988k strongly concordant for relationships of AUTRAN G., DELANOY G. & THOMEL G. (1986).- Dis- ancestor-descendant type between both these cussion critique des genres d'ammonites two groups. déroulées : Toxoceras d'ORBIGNY, Hemiba- According to our hypothesis, the Hemihopli- culites HYATT, 1900 et Moutoniceras SARKAR, tidae first appeared suddenly in the Essaouira- 1955. Proposition d'un nouveau genre Pseu- Agadir Basin at the end of the Early Barremian, domoutoniceras pour l'espèce T. annulare before invading the northern Tethyan margin at d'ORBIGNY.- Comptes Rendus de l'Académie 267 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

des Sciences, Paris, (série II), t. 303, nº 11, horizons pour le Barrémien supérieur.- p. 1059-1064. Carnets de Géologie [Notebooks on BARRON E.J., HARRISON C.G.A., SLOAN J.L. & HAY Geology], Brest, Article 2008/03 W.W. (1981).- Paleogeography, 180 million (CG2008_A03), 18 p. years ago to the present.- Eclogae Geolo- BERT D., DELANOY G. & BERSAC S. (2011).- The gicae Helvetiae, Basel, nº 74, p. 443-470. Dichotomus Horizon: proposal for a new bio- BERT D. (2012).- Phylogenetic relationships chronologic unit of the Giraudi Zone of the among the Hemihoplitidae SPATH, 1924 (Am- Upper Barremian of southeastern France, monoidea, Upper Barremian). In: BERT D. & and considerations regarding the genus Ime- BERSAC S. (eds.), First meeting of the Re- rites ROUCHADZÉ (Ammonoidea, Gassendice- search group for paleobiology and biostrati- ratinae).- Carnets de Géologie [Notebooks graphy of the ammonites – Communica- on Geology], Brest, Article 2011/01 (CG2011_ tions.- Boletin del Instituto de Fisiografia y A01), 12 p. Geologia, Rosario, nº 82, p. 37-38. BERT D., DELANOY G. & CANUT L. (2009).- L'ori- BERT D. (2013).- Factors of intraspecific varia- gine des Imerites ROUCHADZÉ, 1933 : résultat bility in ammonites, the example of Gas- d'une innovation chez les Gassendiceratinae sendiceras alpinum (d'ORBIGNY, 1850) (Hemi- BERT, DELANOY & BERSAC, 2006 (Ammonoidea, hoplitidae, Upper Barremian).- Annales de Ancyloceratina).- Annales de Paléontologie, Paléontologie, Paris, vol. 100, nº 3, p. 217- Paris, vol. 95, p. 21-35. 236. BERT D., BUSNARDO R., DELANOY G. & BERSAC S. BERT D. & BERSAC S. (2012).- Rediscovery of the (2010).- Problems in the identity of "Crioce- type specimen and status of Ezeiceras ras" barremense KILIAN, 1895 (Ancylocera- heberti FALLOT, 1884 (Ammonoidea, Hemi- tida, Late Barremian), and their proposed hoplitidae, Barremian). In: BERT D. & BERSAC resolution.- Carnets de Géologie [Notebooks S. (eds.), First meeting of the Research on Geology], Brest, Article 2010/01 group for paleobiology and biostratigraphy of (CG2010_ A01), 17 p. the ammonites – Communications.- Boletin BININDA-EMONDS O.R.P., BRYANT H.N. & RUSSELL del Instituto de Fisiografia y Geologia, Rosa- A.P. (1998).- Supraspecific taxa as terminals rio, nº 82, p. 42-44. in cladistic analysis: implicit assumptions of BERT D. & BERSAC S. (2013).- Evolutionary pat- monophyly and a comparison of methods.- terns – tested with cladistics – and Biological Journal of the Linnean Society, processes in relation to palaeoenvironments London, nº 64, p. 101-133. of the Upper Barremian genus Gassendiceras BODIN S., GODET A., ADATTE T. & FÖLLMI K.B. (Ammonitina, Lower Cretaceous).- Palaeon- (2005).- Palaeoceanographic and palaeo- tology, London, vol. 56, p. 631-646. climatic changes of the northern Tethyan BERT D., BERSAC S., DELANOY G. & CANUT L. realm during the Hauterivian-Barremian: (2013).- Palaeontology, taxonomic revision New insight from the Angles section (SE and variability of some species of the genus France).- Géologie Alpine, (série "Colloques Gassendiceras BERT et al., 2006 (Ammo- et excursions"), Grenoble, nº 7, p. 138-146. nitina, Upper Barremian) from southeastern BODIN S., FIET N., GODET A., MATERA V., WESTER- France.- Acta Geologica Polonica, Warsaw, MANN S., CLÉMENT A., JANSSEN N.M.M., STILLE vol. 63, p. 355-397. P. & FÖLLMI K.B. (2009).- Early Cretaceous BERT D. & DELANOY G. (2000).- Considérations (late Berriasian to early Aptian) palaeocea- nouvelles sur quelques représentants bar- nographic change along the northwestern rémiens des Puchelliidae DOUVILLÉ, 1890 et Tethyan margin (Vocontian Trough, southe- 13 18 des Hemihoplitidae SPATH, 1924 (Ammonoi- astern France): δ C, δ O and Sr-isotope dea).- Annales du Muséum d'Histoire Natu- belemnite and whole-rock records.- Creta- relle de Nice, t. XV, p. 63-89. ceous Research, London, vol. 30, p. 1247- BERT D. & DELANOY G. (2009).- Pseudosha- 1262. sticrioceras bersaci nov. sp. (Ammonoidea, BREMER K. (1994).- Branch support and tree sta- Gassendiceratinae), and new ammonite bio- bility.- Cladistics, New York, vol. 10, p. 295– horizon for the Upper Barremian of south- 304. eastern France.- Carnets de Géologie [Note- BRYANT H.N. (1997).- Hypothetical ancestors books on Geology], Brest, Article 2009/02 and rooting in cladistic analysis.- Cladistics, (CG2009_A02), 22 p. New York, vol. 13, p. 337-348. BERT D., DELANOY G. & BERSAC S. (2006).- De- COMPANY M., SANDOVAL J., TAVERA J.M., AOUTEM M. scriptions de représentants nouveaux ou peu & ETTACHFINI M. (2008).- Barremian ammo- connus de la Famille des Hemihoplitidae nite faunas from the western High Atlas, SPATH, 1924 (Barrémien supérieur, Sud-Est Morocco – Biostratigraphy and palaeobiogeo- de la France) : conséquence taxinomique et graphy.- Cretaceous Research, London, vol. phylétiques.- Annales du Muséum d'Histoire 29, nº 1, p. 9-26. Naturelle de Nice, t. XXI, p. 179-253. DELANOY G. (1992).- Les ammonites du Barré- BERT D., DELANOY G. & BERSAC S. (2008).- Propo- mien supérieur de Saint-Laurent de l'Esca- sitions pour un nouveau découpage bio- rène (Alpes-Maritimes, Sud-Est de la Fran- zonal ammonitique, et nouveaux bio- ce).- Annales du Muséum d'Histoire Naturelle

268 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

de Nice, t. IX, 148 p. groups as natural units of biological classi- DELANOY G. (1994).- Les biozones à Feraudia- fication.- Taxon, Chambésy, nº 59, nº 6, p. nus, Giraudi et Sarasini du Barrémien supé- 1641-1653. rieur de la région stratotypique d'Angles- IMMEL H. (1978).- Die Crioceratiten (Ancylocera- Barrême-Castellane (Sud-Est de la France).- tina, Ammonoidea) des mediterranen und Géologie Alpine, Grenoble, (Mémoire H.S.), Borealen Hauterive-Barreme (Unterkreide).- nº 20, p. 279-319. Palaeontographica A, Stuttgart, vol. 163, nº DELANOY G. & FÉRAUD P. (1995).- On the Genus 1-3, 85 p. Spinocrioceras KEMPER, 1973 (Ammonoidea, IMMEL H. (1979).- Über den Ursprung der Bo- Ancyloceratina).- Geologisches Jahrbuch, realen Crioceratiten und zur Phylogenie der Hannover, (Reihe A), Heft 141, p. 201-223. Gattung Crioceratites Leveillé (Ammonoidea, DELANOY G., MAGNIN A., SÉLÉBRAN M. & SÉLÉBRAN Kreide).- Aspekte der Kreide Europas, Stutt- J. (1991).- Moutoniceras nodosum gart, (IUGS Serie A), nº 6, p. 129-140. d'ORBIGNY, 1850 (Ammonoidea, IMMEL H. & MUTTERLOSE J. (1978).- Barrême-Ce- Ancyloceratina) une très grande ammonite phalopoden aus dem kretasichen Untergrund hétéromorphe du Barrémien inférieur.- des Stadtgebietes von Hannover (N-W- Revue de Paléobiologie, Genève, vol. 10, nº Deutshland).- Paläontologische Zeitschrift, 2, p. 229-245. Stuttgart, vol. 54, p. 241-266. DELANOY G., ROPOLO P., MAGNIN A., AUTRAN G., KAKABADZE M.V. (1981).- The Ancyloceratids of POUPON A. & GONNET R. (1995).- Sur le di- the South of the USSR and their strati- morphisme chez les Ancyloceratina (Ammo- graphic significance.- Trudy geologicheskogo noidea) du Crétacé inférieur.- Comptes Ren- Instituta Akademii Nauk GSSR, nº 71, 221 dus de l'Académie des Sciences, Paris, (Série p. [in Russian]. IIa), t. 321, p. 537-543. KAKABADZE M.V. (1983).- On the Hauterivian- DERCOURT J., GAETANI M., VRIELYNCK B., BARRIER Barremian correlations between the South of E., BIJU-DUVAL B, BRUNET M.F. CADET, J.P., the USSR and certain southern and northern CRASQUIN S. & SANDULESCU M. (eds., 2000).- regions of Europe. In: 2. Symposium Kreide. Atlas Peri-Tethys, Palaeogeographical München 1982.- Zitteliana, München, (Reihe maps.- Commission de la Carte géologique B: Abhandlungen der Bayerischen Staats- du Monde / Commission for the geological sammlung für Paläontologie und Geologie), Map of the World (CCGM/CGMW), Paris, 24 Band 10, p. 501-508. maps, explanatory notes I-XX, 269 p. KAKABADZE M.V. & HOEDEMAEKER P.J. (2004).- FARRIS J.S. (1970).- Methods for computing Heteromorphic ammonites from the Barre- WAGNER trees.- Systematic Zoology, Oxford, mian and Aptian strata of Colombia.- Scripta vol. 19, p. 83–92. Geologica, Leiden, nº 128, p. 39-182. FARRIS J.S. (1989).- The retention index and the KAKABADZE M.V. & HOEDEMAEKER P.J. (2010).- rescaled consistency index.- Cladistics, New New data on Early Cretaceous (Hauterivian- York, vol. 5, nº 4, p. 417-419. Barremian) heteromorphic ammonites from GOLOBOFF P.A., FARRIS J.S. & NIXON K.C. (2008).- northern Germany.- Scripta Geologica, TNT, a free program for phylogenetic ana- Leiden, nº 140, 168 p. lysis.- Cladistics, New York, vol. 24, nº 5, p. KAKABADZE M.V. & THIEULOY J.-P. (1991).- Am- 774–786. monites hétéromorphes du Barrémien et de GUEX J. (2006).- Reinitialization of evolutionary l'Aptien de Colombie (Amérique du Sud).- clocks during sublethal environmental stress Géologie Alpine, Grenoble, nº 67, p. 81-113. in some invertebrates.- Earth and Planetary KLEIN J., BUSNARDO R., COMPANY M., DELANOY G., Science Letters, La Jolla, vol. 242, p. 240– KAKABADZE M., REBOULET S., ROPOLO P., VASI- 253. CEK Z. & VERMEULEN J. (2007).- Lower Creta- HARDENBOL J., THIERRY J., FARLEY M.B., JACQUIN T., ceous Ammonites III Bochianitidae, Protan- GRACIANSKY P.C. de & VAIL P.R. (1998).- Me- cyloceratoidea, Ancyloceratoidea, Ptycho- sozoic and Cenozoic sequence chronostrati- ceratoidea. In: RIEGRAF W. (ed.), Fossilium graphic framework of European Basins. In: Catalogus I: Animalia.- Backhuys Publishers, GRACIANSKY P.C. de, HARDENBOL J., JACQUIN T. Leiden, 381 p. & VAIL P.R. (eds.), Mesozoic and Cenozoic KLINGER H.C. & KENNEDY W.J. (1992).- Creta- sequence stratigraphy of European basins.- ceous faunas from Zululand and Natal, Society of Economic Paleontologists and South Africa. Barremian representatives of Mineralogists, Special Publication, Tulsa, nº the ammonite family Ancyloceratidae GILL, 60, p. 3-13. 1871.- Annals of the South African Museum, HÖRANDL E. (2006).- Paraphyletic versus mono- Cape Town, vol. 101, p. 71-138. phyletic taxa - evolutionary versus cladistic KLUGE A.G. & FARRIS J.S. (1969).- Quantitative classifications.- Taxon, Chambésy, vol. 55, phyletics and the evolution of anurans.- nº 55, p. 564-570. Systematic Zoology, Oxford, vol. 18, nº 1, p. HÖRANDL E. (2007).- Neglecting evolution is bad 1-32. taxonomy.- Taxon, Chambésy, vol. 56, nº 1, KORNET D.J. & TURNER H. (1999).- Coding poly- p. 1-5. morphism for phylogeny reconstruction.- HÖRANDL E. & STUESSY T.F. (2010).- Paraphyletic Systematic Biology, Oxford, vol. 48, p. 365-

269 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

379. RAWSON P.F. (1995).- The "Boreal" Early Creta- KRON K. A. & JUDD W.S. (1990).- Phylogenetic ceous (pre-Aptian) ammonite sequences of relationships within the Rhodoreae (Erica- NW Europe and their correlation with the ceae) with specific comments on the pla-ce- western Mediterranean faunas.- Memorie de- ment of Ledum.- Systematic Botany, Lara- scrittive della Carta geologica d'Italia, Roma, mie, vol. 15, p. 57-68. vol. 51, p. 121-130. MCARTHUR J.M., JANSSEN N.M.M., REBOULET S., REBOULET S. (2007).- Diversification des am- LENG M.J., THIRLWALL M.F. & SCHOOTBRUGGE B. monoïdes hétéromorphes : l'exemple des van de (2007).- Palaeotemperatures, polar Himantoceras. Evolution et changements ice-volume, and isotope stratigraphy paléoenvironnementaux au Valanginien.- (Mg/Ca, δ18O, δ13C, 87Sr/86Sr): The Early Bulletin de l'Association géologique Auboise, Cretaceous (Berriasian, Valanginian, Troyes, nº 28, p. 13-35. Hauterivian).- Palæogeography, Palæoclima- REBOULET S. & ATROPS F. (1995).- Rôle du climat tology, Palæoecology, Amsterdam, vol. 248, sur les migrations et la composition des peu- n° 3-4, p. 391-430. plements d'ammonites du Valanginien supé- MAYR E. (1974).- Populations, espèces et évo- rieur du bassin vocontien (S-E de la Fran- lution.- Hermann, Paris, 496 p. ce).- Geobios, Villeurbanne, vol. 28, Supple- MIKHAILOVA I. A. & BARABOSHKIN E.Yu. (2009).- ment 1, p. 357-365. The evolution of the heteromorph and mono- REBOULET S., ATROPS F., FERRY S. & SCHAAF A. morph Early Cretaceous Ammonites of the (1992).- Renouvellement des ammonites en Suborder Ancyloceratina WIEDMANN.- Paleon- fosse vocontienne à la limite Valanginien- tological Journal, nº 43, p. 527-536. Hauterivien.- Geobios, Villeurbanne, vol. 25, MONNET C. (2005).- Anisian (Middle Triassic) fasc. 4, p. 469-476. and Cenomanian (mid-Cretaceous) ROCH E. (1920).- Études géologiques dans la ré- Ammonoids: biochronology, biodiversity, gion méridionale du Maroc occidental.- Notes and evolution trends.- Ph.D. Thesis, et Mémoires du Service de la Carte géologi- University of Zürich, 706 p. que du Maroc, Rabat 542 p. MUTTERLOSE J. & BÖCKEL B. (1998).- The Barre- ROPOLO P. & GONNET R. (1995).- Nouveaux mian - Aptian interval in NW Germany: a re- exemples de dimorphisme chez les Ancy- view.- Cretaceous Research, London, vol. loceratina (Ammonoidea) de l'Hauterivien 19, p. 539-568. vocontien.- Géologie Méditerranéenne, MUTTERLOSE J. & BORNEMANN A. (2000).- Distribu- Marseille, t. XXII, p. 93-109. tion and facies patterns of Lower Cretaceous ROUGET I., NEIGE P. & DOMMERGUES J.L. (2004).- sediments in northern Germany: a review.- L'analyse phylogénétique chez les ammoni- Cretaceous Research, London, vol. 21, p. tes : état des lieux et perspectives.- Bulletin 733-759. de la Société géologique de France, Paris, NEIGE P., ROUGET I. & MOYNE S. (2007).- vol. 175, p. 507-512. Phylogenetic practices among scholars of SARKAR S.S. (1955).- Révision des Ammonites fossil cephalopods, with special reference to déroulées du Crétacé inférieur du SE de la cladistics. In: LANDMAN N.H., DAVIS R.A. & France.- Mémoires de la Société géologique MAPES R.H. (eds.), Cephalopods present and de France, Paris, (N.S.), t. XXXIV, nº 72, past: new insights and fresh perspectives.- 176 p. Springer-Verlag Inc., New York, p. 3-14. SIMMONS N.B. & GLEISER J.H. (2002).- Sensitivity NIXON K.C. & DAVIS J.I. (1991).- Polymorphic analysis of different methods of coding taxo- taxa, missing values and cladistic analysis.- nomic polymorphism: an example from Cladistics, New York, vol. 7, nº 3, p. 233- higher-level bat phylogeny.- Cladistics, New 241. York, nº 18, p. 571-584. OGG J.G. & HINNOV L.A. (2012).- Cretaceous, SPATH L.F. (1924).- On the ammonites of the Chapter 27. In: GRADSTEIN F.M., OGG J.G., Speeton Clay and the subdivisions of the SCHMITZ M. & OGG G. (eds.), The Geologic Neocomian.- Geological Magazine, Cambrid- Time Scale 2012 2-Volume Set.- Elsevier, ge, vol. 61, nº 2, p. 73-89. Amsterdam, p. 793-853. SWOFFORD D.L. & MADDISON W.P. (1987).- Re- PRENDINI L. (2001).- Species or supraspecic taxa constructing ancestral character states under as terminals in cladistic analysis? Ground- WAGNER parsimony.- Mathematical Bio- plans versus exemplars revisited.- Syste- sciences, Atlanta, vol. 87, p. 199–229. matic Biology, Oxford, vol. 50, p. 290-300. SWOFFORD D.L. & MADDISON W.P. (1992).- Parsi- RAWSON P.F. (1975).- The interpretation of the mony, character-state reconstructions, and Lower Cretaceous heteromorph ammonite evolutionary inferences. In: MAYDEN R.L. genera Paracrioceras and Hoplocrioceras (ed.), Systematics, historical ecology, and SPATH, 1924.- Palaeontology, London, vol. North American freshwater fishes.- Stand- 18, p. 275-283. ford University Press, p. 186–223. RAWSON P.F. (1983).- The Valanginian to Aptian VERMEULEN J. (1995).- Nouvelle biozonation ba- stages – current definitions and outstanding sée sur la famille des Pulchelliidae (Ammo- problems.- Zitteliana, Munich, Band 10, p. noidea).- Géologie Alpine, Grenoble, nº 71, 493-500. p. 199-211.

270 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

VERMEULEN J. (1997).- Moutoniceras marii, nou- se), Jahrgang 1962, nº 1, 148 p. velle espèce d'ammonite hétéromorphe du WISSLER L., WEISSERT H., MASSE J.-P. & BULOT L. Barrémien du Sud-Est de la France.- Riviéra (2002).- Chemostratigraphic correlation of Scientifique, Nice, p. 73-80. Barremian and lower Aptian ammonite zones VERMEULEN J. (2000).- Nouvelles données sur les and magnetic reversals.- International Jour- répartitions stratigraphiques, les évolutions nal of Earth Science (Geologische Rund- et les classifications de trois familles d'am- schau), Mendig, vol. 91, nº 2, p. 272–279. monites du Crétacé inférieur.- Géologie Alpi- WRIGHT C.W., CALLOMON J.H. & HOWARTH M.K. ne, Grenoble, nº 75, p. 123-132. (eds., 1996).- Cretaceous Ammonoidea. In: WIEDMANN J. (1962).- Underkreide-Ammoniten KAESLER R.L. (ed.), Mollusca 4 (revised).- von Mallorca. 1. Lieferung: Lytoceratina, Ap- Treatise on Invertebrate Paleontology, Geo- tychi.- Abhandlungen der Akademie der Wis- logical Society of America, New York; Uni- senschaften und der Literatur, Mayence, versity of Kansas, Lawrence, Part L, 362 p. (Mathematisch-naturwissenschaftliche Klas-

271 Carnets de Géologie [Notebooks on Geology] - vol. 14, n° 13

Appendix List of characters 00. Adult size: (0) little; (1) median; (2) large; (3) very large. 01. Coiling shape: (0) evolute normally coiled; (1) crioconic uncoiled; (2) tripartite tight; (3) tripartite strict. 02. Height whorl growth: (0) low; (1) moderate; (2) high. 03. Whorl section shape: (0) thick; (1) compressed. 04. Coiling hiatus: (0) almost joined whorls; (1) weak; (2) large. 05. Heberti stage: (0) absent; (1) present. 06. Barremense stage: (0) absent; (1) present. 07. Spines: (0) absent; (1) long and thin; (2) short and massive. 08. Shape of the main ribs on the venter: (0) not or mainly not altered; (1) attenuated. 09. Looped ribs: (0) absent; (1) present. 10. Smooth interribs: (0) rare; (1) frequent. 11. Bifurcated or polyfurcated ribs: (0) absent or scarce; (1) present. 12. Fibulate ribs: (0) absent; (1) present. 13. Wedge-shaped ribs: (0) absent; (1) present. 14. Position of lateral tubercles: (0) absent; (1) upper third; (2) upper quarter. 15. Shape of the tubercles: (0) slender; (1) robust. 16. Simplified ornamentation stage: (0) absent; (1) present. 17. Shape of the flanks: (0) flattened; (1) rounded. 18. Shape of the dorsum: (0) flattened; (1) concave; (2) convex.

Taxon-character matrix with generic taxa (generic taxa in white, polymorphic states in red):

Taxa / Characters 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 Crioceratites nolani 2 1 0 [01] 2 0 0 1 0 0 1 0 0 0 1 0 0 0 0 Emericiceras 3 2 0 [01] 1 0 0 1 0 1 1 1 1 0 1 0 0 [01] 0 Fissicostaticeras 2 1 1 [?] 1 0 [01] 2 0 1 1 1 [01] 0 2 [01] 1 0 0 Gassendiceras essaouirae nov. sp. [?] 1 1 0 1 1 1 2 1 [?] 0 0 0 1 1 1 1 1 0 G. multicostatum 2 1 1 0 1 1 1 2 1 1 0 0 0 1 1 1 1 1 0 Honnoratia 3 2 0 [01] 1 0 0 1 0 1 1 0 1 0 1 0 0 1 2 Moutoniceras 3 3 0 [01] 2 0 0 0 1 0 [01] 0 [01] 1 0 [01] 0 0 2 Paracrioceras stadtlaenderi 2 1 1 0 1 [?] 1 2 0 1 1 1 0 0 2 1 1 1 0 Pseudothurmannia [01] 0 2 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 Sornayites 1 0 1 [01] 0 0 0 2 0 0 1 1 0 0 1 1 0 [01] 1 Toxancyloceras 2 3 0 1 2 0 0 1 [01] 0 1 0 [01] 1 1 1 0 1 2

Modified taxon-character matrix with generic taxa replaced by species: Taxa / Characters 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18 Crioceratites nolani 2 1 0 [01] 2 0 0 1 0 0 1 0 0 0 1 0 0 0 0 Emericiceras emerici 3 2 0 [01] 1 0 0 1 0 1 1 1 1 0 1 0 0 0 0 Fissicostaticeras fissicostatum 2 1 1 [?] 1 0 1 2 0 1 1 1 [01] 0 2 1 1 0 0 Gassendiceras essaouirae nov. sp. [?] 1 1 0 1 1 1 2 1 [?] 0 0 0 1 1 1 1 1 0 G. multicostatum 2 1 1 0 1 1 1 2 1 1 0 0 0 1 1 1 1 1 0 Honnoratia thiollerei 3 2 0 [01] 1 0 0 1 0 1 1 0 1 0 1 0 0 1 2 Moutoniceras moutonianum 3 3 0 [01] 2 0 0 0 1 0 1 0 [01] 1 0 1 0 0 2 Paracrioceras stadtlaenderi 2 1 1 0 1 [?] 1 2 1 1 1 0 0 0 2 1 1 1 0 Pseudothurmannia picteti 1 0 2 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 Sornayites paronai 1 0 1 [01] 0 0 0 1 0 0 1 1 0 0 1 1 0 0 1 Toxancyloceras vandenheckei 2 3 0 1 2 0 0 1 1 0 1 0 [01] 1 1 1 0 1 2

272