This is the accepted version of the article: Matamales Andreu, Rafel; Juárez Ruiz, Josep; Pictet, Antoine; [et al.]. «Glauco- ceras gen. nov., a small uncoiled ammonoid from the Tethyan late Hauterivian (Early Cretaceous) : evolutionary implications at the dawn of the diversification of heteromorphic lineages». Cretaceous research, Vol. 100 (August 2019), p. 114-125. DOI 10.1016/j.cretres.2019.02.025 This version is avaible at https://ddd.uab.cat/record/215295 under the terms of the license Accepted Manuscript Glaucoceras gen. nov., a small uncoiled ammonoid from the Tethyan late Hauterivian (Early Cretaceous): evolutionary implications at the dawn of the diversification of heteromorphic lineages Rafel Matamales-Andreu, Josep Juárez-Ruiz, Antoine Pictet, Josep Anton Moreno- Bedmar PII: S0195-6671(18)30521-4 DOI: https://doi.org/10.1016/j.cretres.2019.02.025 Reference: YCRES 4099 To appear in: Cretaceous Research Received Date: 20 December 2018 Revised Date: 31 January 2019 Accepted Date: 22 February 2019 Please cite this article as: Matamales-Andreu, R., Juárez-Ruiz, J., Pictet, A., Moreno-Bedmar, J.A., Glaucoceras gen. nov., a small uncoiled ammonoid from the Tethyan late Hauterivian (Early Cretaceous): evolutionary implications at the dawn of the diversification of heteromorphic lineages, Cretaceous Research, https://doi.org/10.1016/j.cretres.2019.02.025. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 Glaucoceras gen. nov., a small uncoiled ammonoid from the Tethyan late 2 Hauterivian (Early Cretaceous): evolutionary implications at the dawn of the 3 diversification of heteromorphic lineages 4 5 Rafel MATAMALES-ANDREU1,2*, Josep JUÁREZ-RUIZ2,3, Antoine PICTET4, Josep PT 6 Anton MORENO-BEDMAR5 RI 7 8 *rafel.matamales@icp.cat (corresponding author). SC 9 1. Institut Català de Paleontologia Miquel Crusafont. Universitat Autònoma de 10 Barcelona, edifici ICTA–ICP, Carrer de les Columnes s/n, Campus de la UAB. 08193, 11 U Cerdanyola del Vallès (Barcelona, Spain). AN 12 2. Museu Balear de Ciències Naturals. Carretera Palma–Port de Sóller Km. 30. 07100, M 13 Sóller (Illes Balears, Spain). 14 3. Societat d’Història Natural de les Balears. Carrer Margalida Xirgu 16, baixos. 07011, D 15 Palma (Illes Balears, Spain). TE 16 4. Musée Cantonal de Géologie, Anthropole. 1015, Lausanne (Vaud, Switzerland). 17 5. Departamento de Paleontología, Instituto de Geología, Universidad Nacional EP 18 Autónoma de México, Ciudad Universitaria, Delegación Coyoacán. 04510, Ciudad de México (Mexico). C 19 AC 20 21 22 Abstract 23 24 The poorly-known species Baculites Renevieri Ooster, 1860 is revised using the type 25 material from Switzerland and new conspecific specimens collected in deposits from ACCEPTED MANUSCRIPT 26 Mallorca and the Betic Cordillera (Western Mediterranean), listing all the previous 27 mentions of the species. The uniqueness of its ornamentation pattern and shell shape 28 compared to other coeval ammonoid faunas demands for the description of Glaucoceras 29 gen. nov. to contain this species. In addition, a thorough comparison of Glaucoceras 30 renevieri to other morphologically reminiscent genera or species is provided. This new PT 31 set of data allows to improve the knowledge of one of the many phylogenetic lines of RI 32 heteromorphic ammonites that diversified over the latest Hauterivian in the 33 Mediterranean Province, also providing hints about the timing of those radiation events. SC 34 This aspect is preliminary explored using data on other coeval heteromorphic 35 ammonoid groups from the literature and new finds, concluding that there occurred two 36 U main radiation events of heteromorphic ammonites at the latest Hauterivian, most likely AN 37 related to environmentally stressful episodes, specifically, transgressive events. M 38 39 Keywords: Mallorca, Betic Cordillera, External Prealps, Ammonoidea, Hauterivian D 40 TE 41 42 1. Introduction EP 43 The Barremian small to medium-sized heteromorphic ammonoids have recently been C 44 AC 45 vastly studied and revised by Vašíček & Wiedmann (1994), Vermeulen (2005, 2006, 46 2009, 2010a, 2010b), Vermeulen & Vašíček (2011) and Vermeulen et al. (2007, 2010a, 47 2012b, 2013a, 2013b, 2014a, 2014b), among others. In the latest Hauterivian (Tab. 1), 48 the variety of forms might be as numerous as in the Barremian (e.g., Vašíček & 49 Hoedemaeker, 2003; Vermeulen et al., 2012a, 2013a; Lukeneder, 2018), however 50 remaining mostly unstudied. In Mallorca and in the Betic Cordillera, the deposits from ACCEPTED MANUSCRIPT 51 the aforementioned age have delivered a great number of forms, belonging both to 52 Protancyloceratina Vermeulen, 2005 or to Ancyloceratina Wiedmann, 1966 (the latter 53 considered herein as a senior synonym of Turrilitina Beznosov & Mikhailova, 1983, 54 sensu Vermeulen, 2006). The present study focuses on a particular genus, Glaucoceras 55 gen. nov., described to contain the poorly known species Baculites Renevieri Ooster, PT 56 1860. All the aforementioned data on this species is compiled, completing its diagnosis RI 57 and comparing it to similar species in order to avoid any possible confusions in their 58 identification. Also, a probable phylogenetical position is suggested based on shell SC 59 shape and ornamental affinities. This enables to clarify the taxonomy of one of the 60 many groups of small heteromorphic ammonites that appeared at the latest Hauterivian. 61 U The better comprehension of these groups will in turn be relevant in future attempts to AN 62 study the evolution of heteromorphic ammonoid shape and lineages through time, most M 63 probably linked to abiotic events, since the heteromorphic groups tended to radiate as a 64 response to environmental stresses (further explored in ‘5. Discussion’ herein). D 65 TE 66 67 2. Geological context and fossil localities EP 68 2.1. External Prealps C 69 AC 70 2.1.1. Veveyse de Châtel (46°31'N 06°55'E) 71 This historical outcrop is located along the Veveyse de Châtel torrent, at about 2 km 72 eastward from the small city of Châtel-St-Denis (Fribourg, Switzerland) (Fig. 1A-a). 73 The stream crosses several geological units such as the Subalpine Molasse and the 74 Préalpes nappes. The external Préalpes nappe has been classically attributed to the 75 Ultrahelvetic domain (Weidmann et al., 1993), a distal hemipelagic setting following ACCEPTED MANUSCRIPT 76 the southern front of the neritic Helvetic domain. The Ultrahelvetic nappe is arranged 77 under the form of a succession of tectonic scales (Gagnebin, 1924; Anatra, 1986; 78 Weidmann et al., 1993). The primary scale, or ‘Riondonnaire scale’, is composed by an 79 isoclinal stratigraphic succession spanning from the Upper Jurassic to the Lower 80 Cretaceous (Gagnebin, 1920). The Veveyse de Châtel section was described in detail by PT 81 several authors (Gagnebin, 1934, 1945; Charollais & Rigassi-Studer, 1961; Rigassi & RI 82 Roveda, 1964), and studied for its palaeontological richness, especially for its 83 ammonoid content by Ooster (1860–1863) and Sarasin & Schöndelmayer (1901–1902). SC 84 A detailed stratigraphic and biostratigraphic work based on ammonites was later carried 85 out by Busnardo et al. (2003) who reviewed the previous existing collections and 86 U collected new material bed-by-bed. They described an excellent profile in the group of AN 87 outcrops located higher up in the torrent, consisting in an alternation of marlstones and M 88 limestones of upper Hauterivian to lowermost Barremian age, which they attributed to 89 their informal ‘Alternances supérieures de marnes et de calcaires tachetés’ unit D 90 (Charollais et al., 1981, 1993). TE 91 92 2.2. Mallorca EP 93 Mallorca, the largest of the Balearic Islands, is included in the Betic System and its series are correlated with the Subbetic Domain (see also ‘2.3. Southeastern Iberian C 94 AC 95 Peninsula’). As for Hauterivian strata, the dominant lithologies consist in marly 96 limestones rhythmically intercalated with marlstones. They are assigned to the 97 ‘Maiolica’ facies, which represent a long-lasting pelagic sedimentation event with 98 particular abundance of ammonites (Colom, 1975, Vera, 2004). The Mallorcan 99 specimens studied in this work come from four different localities (Fig. 1B), all of 100 which have been recently described for the first time by Juárez-Ruiz & Matamales- ACCEPTED MANUSCRIPT 101 Andreu (2015, unpublished). In this work, only the sections of interest within each 102 locality will be cited as follows: 103 104 2.2.1. Biniamar (39°43'N 02°52'E) 105 This outcrop, also mentioned in Bert et al. (2017), is located between the towns of PT 106 Lloseta and (mostly) Biniamar (Fig. 1B-b), and constitutes a nearly complete series RI 107 from the Berriasian to the upper Barremian. However, it is mostly covered by 108 pastureland, thus making it difficult to observe at some points, resulting in several SC 109 stages having only been recognised by ex situ samples. A section comprising three beds 110 from the upper Hauterivian (Krenkeli Subzone, Balearis Zone) has been intensively 111 U sampled (Fig. 1C), delivering three partial specimens of G. renevieri. Previous works on AN 112 Cretaceous outcrops of this area include Fallot (1910, 1922), Fallot & Termier (1923) M 113 and Wiedmann (1962a, 1962b, 1964, 1967), although it is impossible to know whether 114 they referred to the same outcrops we studied as the region is quite rich in fossiliferous D 115 Cretaceous rocks. TE 116 117 2.2.2. Can Negret (39°43'N 02°50'E) EP 118 This quarry, located at the southwest of the town of Lloseta (Fig. 1B-c) and mentioned in Bert et al. (2017), possesses the most complete stratigraphic succession of the Lower C 119 AC 120 Cretaceous of the island of Mallorca, currently under exploitation by the cement 121 industry. Its stratigraphic range covers almost continuously from the Lower Jurassic to 122 the uppermost Lower Cretaceous. However, sections cannot be measured and 123 exhaustively sampled due to the ever-changing nature of the locality. Therefore, the 124 samples are relatively dated upon direct association to index taxa. One fragmentary 125 specimen of G. aff. renevieri has been found in direct association with Balearites ACCEPTED MANUSCRIPT 126 krenkeli (Sarkar, 1955), thus indicating Krenkeli Subzone, Balearis Zone (upper 127 Hauterivian). 128 129 2.2.3. Manacor (39°32'N 03°14'E) 130 This outcrop is located at the south-east of the town of Manacor (Fig. 1B-d), near the PT 131 road Ma-4015. In situ stratigraphic sections from the Binelli Subzone to the Picteti RI 132 Subzone (upper Hauterivian) have been recorded, although they are unfortunately 133 mostly covered by pastureland and therefore most of the samples were recovered from SC 134 ex situ rocks. A fragment of G. renevieri was found in the Krenkeli Subzone, in 135 association with Balearites ‘majoricensis’ (Nolan, 1894), while the only probable 136 U subadult part has been found related to the index species of the aforementioned subzone. AN 137 M 138 2.2.4. Son Macià (39°30'N 03°13'E) 139 The fields at the north and west of the village of Son Macià (Fig. 1B-e), used as D 140 pastureland, contain several sections exposing upper Hauterivian rocks from the Ligatus TE 141 Zone to the Mortilleti Subzone (Ohmi Zone). In an ex situ rock, however in association 142 with a fragment of Pseudothurmannia pseudomalbosi Sarasin & Schöndelmayer, 1901 EP 143 (exclusive from the P. mortilleti Subzone, as concluded by Company et al., 2003), one partial specimen of G. renevieri has been recovered. C 144 AC 145 146 147 2.3. Southeastern Iberian Peninsula 148 2.3.1. Arroyo Gilico (38°09'N 01°40'W) 149 The Arroyo Gilico section (Murcia, SE Spain) (Fig. 1D-f) is part of the External 150 Subbetic, in the Betic External Zones, within the Betic Cordillera. The Betic System ACCEPTED MANUSCRIPT 151 constitutes a strip of variable width, broadly arranged following a SW–NE orientation. 152 Its sediments are variably represented depending on the zone, combining episodes with 153 high sedimentation rates, as well as condensed sequences and even stratigraphic 154 hiatuses, mainly caused by the tectonic activity (Vera, 2004). In the zone referred in the 155 present work, the Cretaceous is mostly represented by pelagic sequences (see Vera, PT 156 2004; Company et al., 2005). A detailed biostratigraphic approach on the Arroyo Gilico RI 157 section has been recently carried out (Aguado et al., 2008, 2014), although some fossils 158 of this locality have been studied in other works (e.g., Company et al., 2003, 2005, SC 159 2010; Vermeulen et al., 2010b; Matamales-Andreu, 2017a, 2017b; Matamales-Andreu 160 & Company, 2018). The outcrop has been extensively bed-by-bed sampled over the 161 U years by different researchers. This section covers from the Balearis Zone (upper AN 162 Hauterivian) to the Vandenheckei Zone (upper Barremian) (Aguado et al., 2008). For M 163 detailed stratigraphic description see Aguado et al. (2014). 164 D 165 TE 166 3. Material and methods 167 EP 168 For this work, a total of 13 fragmentary specimens have been studied. The type specimen of Baculites renevieri Ooster is housed in the Naturhistorisches Museum Bern C 169 AC 170 (NMBE) (Bern, Switzerland). The specimens from Mallorca are deposited in the Museu 171 Balear de Ciències Naturals (MBCN) (Sóller, Balearic Islands, Spain). The ones from 172 the Arroyo Gilico section are part of the Company-Sandoval-Tavera collection, housed 173 in the Universidad de Granada (X.V1 refers to the Arroyo Gilico section) (Granada, 174 Andalucía, Spain). All the specimens have been measured using a manual calliper or by 175 digital means on the scaled pictures. The measurements have been taken as follows: [1] ACCEPTED MANUSCRIPT 176 The minimum and maximum shell width in both preserved ends of the shell, [2] The 177 maximum preserved length of the shell, [3] The growth rate calculated as ((maximum 178 shell width) – (minimum shell width)) : (shell length), [4] The number of constrictions 179 in the preserved fragment of the shell, [5] The number of constrictions per centimetre, 180 calculated as (number of constrictions) : (maximum length). The terminology for the PT 181 regions of heteromorphic tripartite shells has been taken from Vašíček (1972), that is, RI 182 proversum for the first shaft, flexus for the bend and retroversum for the second shaft. 183 The biostratigraphic framework follows Matamales-Andreu & Company (2018), which SC 184 is slightly modified from the standard zonation presented by Reboulet et al. (2018). 185 186 U AN 187 4. Systematic palaeontology M 188 189 Order: Ammonitida Agassiz, 1847 D 190 Suborder: Protancyloceratina Vermeulen, 2005 TE 191 Superfamily: Bochianitoidea Spath, 1922 192 Family Bochianitidae Spath, 1922 EP 193 Genus Glaucoceras gen. nov. C 194 AC 195 Type species: Baculites Renevieri Ooster, 1860 (p. 91, pl. 60, fig. 4). 196 197 Derivation of name: Because of the morphological similarity of the constantly 198 constricted shell of the ammonite to the segmented and straight loments of the extant 199 Coronilla valentina subsp. glauca, a bush of the subfamily Papilionoideae, widespread 200 over the calcareous soils of the Mediterranean floristic region. ACCEPTED MANUSCRIPT 201 202 Diagnosis: Medium size, probably up to about 10 cm. Heteromorphic shell shape, most 203 probably baculiconic, although a hamuliconic or ptychoconic shape cannot be definitely 204 ruled out. Subcircular section; flanks, dorsum and venter convex. Width growth rate 205 descending over ontogeny. Ornamentation pattern consisting in more or less sharp, PT 206 slightly prosriradiate constrictions that may result in very wide, rounded ribs. They RI 207 cross the venter without alteration, and become less marked on the dorsum. 208 SC 209 Specific content: Glaucoceras renevieri (Ooster, 1860). 210 211 U Stratigraphic and geographic distribution: Latest Hauterivian, from the B. binelli AN 212 Subzone (B. balearis Zone) to the P. mortilleti Subzone (P. ohmi Zone). Reported from M 213 Switzerland (Ooster, 1860), southern Iberian Peninsula and the Balearic Islands (the 214 present work) (Tab. 1). D 215 TE 216 Remarks: Glaucoceras is similar to some genera of Protancyloceratina, especially in 217 their initial stages. Glaucoceras is reminiscent of Bochianites Lory, 1898, but they can EP 218 be differentiated because the latter may either bear attenuated prosriradiate ribs or simply lack any kind of ornamentation, in contrast of the annular constrictions of C 219 AC 220 Glaucoceras. Some species of Bochianites can also develop constrictions, but are 221 accompanied by ribs, and are never as deep and conspicuous as the ones in Glaucoceras 222 (see Mandov, 1971). Finally, the genus Bochianites is thought to disappear in the 223 earliest Hauterivian (e.g., Company, 1987; Reboulet, 1996), whereas Glaucoceras has 224 only been found in the latest Hauterivian. Baculina d’Orbigny, 1850 sensu Vašíček 225 (1999) is similar to Glaucoceras in its baculiconic shape and ornamentation based on ACCEPTED MANUSCRIPT 226 sporadic, well-marked, straight and prosriradiate constrictions. The main difference is 227 that the constrictions of Baculina rouyana d’Orbigny, 1850 sensu Vašíček (1999) are 228 more spaced, and it bears wide, inconspicuous intercalated ribs. In the diagnosis of the 229 holotype of that species, however, neither d’Orbigny (1850) nor Cottreau (1934) ever 230 mentioned the presence of constrictions, so Vašíček (1999)’s specimen could belong to PT 231 a different species altogether. Nevertheless, the stratigraphic range of both specimens RI 232 appears to be restricted to the Valanginian (Cottreau, 1934; Vašíček, 1999), contrarily to 233 Glaucoceras, which appears exclusively in the latest Hauterivian. Glaucoceras is SC 234 similar to Hamulinites Paquier, 1900, with which it shares part of its stratigraphic range, 235 especially in the ornamentation of the initial ontogenetic stages of the latter. However, 236 U they may be differentiated on the basis of stronger, denser, thinner and sharper AN 237 costulation of Hamulinites. Also, Glaucoceras appears to be larger in size. The juvenile M 238 stage of Euptychoceras Breistroffer, 1952 possesses similar dimensions and 239 morphology than Glaucoceras, but they may be differentiated because the former does D 240 not possess any kind of ornamentation on the juvenile region, only developing it further TE 241 in the ontogeny, consisting in very faint, wide, weakly tuberculated ribs. 242 EP 243 There are also some important similarities of Glaucoceras and the juvenile stages of a several genera of Anahamulinidae Breistroffer, 1952, sharing the baculiconic initial C 244 AC 245 shell shape (which later becomes hamuliconic in the Anahamulinidae), even though 246 their ornamentation pattern is always different. In this sense, Amorina Vermeulen, 2005 247 and Auritina Egoian, 1989 differ from Glaucoceras in their thinner and denser 248 ornamentation based on prosriradiate ribs that may or may not be accompanied by 249 sporadic constrictions (see Supplementary Material 1), as well as because of the larger 250 size of the adult specimens of the former genera. Anahamulina Hyatt, 1900, Vasicekina ACCEPTED MANUSCRIPT 251 Vermeulen, 2005, Lazarina Vermeulen, 2009 and Mascarellina Vermeulen, 2009 are 252 more similar in size, but their ornamentation still consists in thin, sharp and 253 prosriradiate ribs, in contrast with the constrictions of Glaucoceras. This same problem 254 occurs with genera such as Ptychohamulina Vermeulen, 2005, Leptohamulina 255 Vermeulen, 2006, Guiomarina Vermeulen et al., 2007, Bulotina Vermeulen, 2009, PT 256 Davouxina Vermeulen, 2009, Badina Vermeulen & Vašíček, 2011 and Defayella RI 257 Vermeulen, 2013b that, apart from the thinner and sharper costulation on the 258 proversum, possess a smaller adult size. Similarly, Djamaina Vermeulen et al., 2010a, SC 259 Baqueina Vermeulen et al., 2010a, Pacaudina Vermeulen et al., 2010a and Curiolina 260 Vermeulen, 2010a are smaller than Glaucoceras in their adult size, but this group does 261 U possess thicker ribs, that in some cases might remind of the interspaces between AN 262 constrictions of Glaucoceras. They may be differentiated because the ribs are more M 263 prosriradiate and always sharper in the former genera group, never forming true 264 constrictions. Duyeina Vermeulen, 2005 is perhaps the most similar genus, both in D 265 terms of shell size and ornamentation. It possesses a pattern based on straight, simple TE 266 and very thick, slightly prosriradiate ribs, which could be confused with the interspaces 267 between constrictions of Glaucoceras. However, the interspaces between ribs in EP 268 Duyeina are clearly not as sharp as the constrictions of Glaucoceras. What is more, their stratigraphic range is quite distant: latest Hauterivian for Glaucoceras opposed to late C 269 AC 270 Barremian for Duyeina spp. (Vermeulen, 2005), making homeomorphy the most 271 plausible explanation for their morphological coincidence. Finally, their phylogenetic 272 relationships seem to be rather unlinked as well, since Duyeina probably evolved from 273 some Barremian Anahamulinidae such as Pacaudina Vermeulen et al., 2010a 274 (Vermeulen et al., 2012b). Glaucoceras, on the other hand, is herein thought to be 275 related to the late Hauterivian Euptychoceras (see ‘Remarks’ below). In fact, the first ACCEPTED MANUSCRIPT 276 occurrences of all of the mentioned genera in this paragraph are younger than the last 277 occurrence of Glaucoceras (Barremian vs. Hauterivian) except for Amorina and 278 Anahamulina, which are coeval. 279 280 Glaucoceras also shares a similar morphology to many Baculitidae Gill, 1871, although PT 281 this group is much younger. It differs from Baculites Lamarck, 1799 because the latter RI 282 does not bear any kind of ornamentation on juvenile stages (Glaucoceras already 283 possesses the constrictions) and Lechites Nowak, 1908, because it only possesses faint SC 284 constrictions or very weak costulation on its juvenile stage. The similarity is greater in 285 the case of Sciponoceras Hyatt, 1894, since this genus is characterised by an 286 U ornamentation pattern consisting in deep and sharp constrictions. However, bearing in AN 287 mind that this genus is stratigraphically very distant (Hauterivian vs. Albian– M 288 Cenomanian), and that it is a direct descendant of Lechites (see Monks, 1999), any 289 relationship with Glaucoceras besides homeomorphy seems unlikely. D 290 TE 291 Glaucoceras gen. nov. is herein provisionally referred to the family Bochianitidae 292 Spath, 1922. Nevertheless, and due to lack of well-preserved suture information, only EP 293 shell morphology and stratigraphic position can be taken into account to support the hypotheses presented in this work. Any relationship with Anahamulinidae Breistroffer, C 294 AC 295 1952 (in the suborder Ancyloceratina Wiedmann, 1966, superfamily Hamulinoidea Gill, 296 1871, family Anahamulinidae Breistroffer, 1952) has been dismissed because the latter 297 possess a hamuliconic shell shape, with slightly curved initial stages, and a juvenile 298 ornamentation pattern consisting in very thin, dense, prosriradiate ribs. Moreover, the 299 first Anahamulinidae appeared in the B. binelli Subzone, at the same time that 300 Glaucoceras. Whereas the former preserved the ornamentation pattern of their supposed ACCEPTED MANUSCRIPT 301 ancestors (Megacrioceratidae Vermeulen, 2006), Glaucoceras shows a deeply modified 302 ornamentation based on constrictions without any trace of ribs. Similar reasons make it 303 possible to exclude Glaucoceras from the Leptoceratoididae Thieuloy, 1966 (in the 304 suborder Protancyloceratina Vermeulen, 2005): the shell shape of the latter may range 305 from hamuliconic, to toxoconic, to more or less crioconic, albeit never baculiconic. PT 306 Moreover, the ornamentation pattern of the Leptoceratoididae consists in thin, dense, RI 307 prosriradiate ribs, which may bifurcate or bear tubercles. Bochianitidae is the only 308 phylogenetic group that fully agrees with the morphological characters of Glaucoceras. SC 309 Although the only coeval representative of this family is Euptychoceras, which 310 possesses no sensible ornamentation on the proversum and has a ptychoconic shell 311 U shape, Glaucoceras is reminiscent of several older Bochianitidae, which are in turn the AN 312 supposed ancestors of Euptychoceras. Therefore, Glaucoceras may be considered as a M 313 late Hauterivian offshoot of Euptychoceras, or, however less likely, it may represent a 314 surviving member of the Bochianites lineage beyond the basal Hauterivian. D 315 TE 316 317 Glaucoceras renevieri (Ooster, 1860) EP 318 (Fig. 2A–K) v. pars *1860 Baculites Renevieri Ooster 1860; Ooster, p. 91, pl. 60, fig. 4, non fig. 5 C 319 AC 320 (=Amorina pictetiformis). 321 1902 Bochianites Renevieri, Ooster; Sarasin & Schöndelmayer, p. 180. 322 1952 Bochianites (?) Renevieri Oost. 1860 sp.; Breistroffer, p. 51. 323 non ? 1979 Bochianites cf. renevieri (Ooster, 1860); Klinger & Kennedy, p. 17, fig. 3, E 324 (=Bochianites sp.). ACCEPTED MANUSCRIPT 325 ? 2001 Bochianites cf. renevieri (OOSTER); Avram, p. 66, pl. 1, fig. 3 (=Bochianites? 326 sp.). 327 2007 Bochianites? renevieri (Ooster, 1860); Klein et al., p. 7. 328 2017 Bochianitinae indet.; Tajika et al., p. 37, fig. 9, AH. 329 PT 330 Type specimens: Syntypes depicted by Ooster, 1860, pl. 60, fig. 4, 5. We hereby RI 331 establish the specimen NMBE 5007731, illustrated by Ooster (1860: pl. 60, fig. 4) as 332 the lectotype, because it undoubtedly possesses the main diagnostic characters of this SC 333 species (herein depicted in Fig. 2A). 334 335 U Studied material: Eleven fragmentary specimens; one from an undetermined AN 336 Hauterivian? bed of the Veveyse de Châtel (lectotype, NMBE 5007731), one from the M 337 Binelli Subzone of Arroyo Gilico (X.V1(–16)9), three from the Krenkeli Subzone of 338 Biniamar (MBCN 23392, MBCN 23394, MBCN 23395), two from the Krenkeli D 339 Subzone of Manacor (MBCN 23390, MBCN 23391), one from the Krenkeli Subzone of TE 340 Arroyo Gilico (X.V1(–10)21), two from the Angulicostatus Subzone of Arroyo Gilico 341 (X.V1(–1)17, X.V1(–2)57) and one from the Mortilleti Subzone of Son Macià (MBCN EP 342 23389). C 343 AC 344 Stratigraphic and geographic distribution: Latest Hauterivian, from the B. binelli 345 Subzone (B. balearis Zone) to the P. mortilleti Subzone (P. ohmi Zone). Reported from 346 Switzerland (Ooster, 1860) and Spain (the present work). 347 348 Emended diagnosis: Medium size, which may reach about 10 cm. Heteromorphic shell 349 shape, most probably baculiconic, although a hamuliconic or ptychoconic shape cannot ACCEPTED MANUSCRIPT 350 be entirely ruled out. Subcircular section; flanks, dorsum and venter convex. Very 351 simple ornamentation consisting in deep and sharp, slightly prosriradiate, straight 352 constrictions, with progressively larger interspaces. They cross the venter without 353 alteration and are very slightly weakened on the dorsum. Suture line unknown. 354 PT 355 Ontogenetic and intraspecific variation: Within the pool of Swiss, Mallorcan and RI 356 Murcian specimens, certain characters, such as the separation of the constrictions, have 357 been deemed slightly variable in a same ontogenetic stage (e.g., compare Fig. 2A with SC 358 Fig. 2C). Likewise, separation of constrictions tends to increase along the ontogeny 359 (e.g., Fig. 2A), although some other specimens in later growth stages seem to refute this 360 U hypothesis (Fig. 2J). Ontogenetic variation of whorl width growth has also been AN 361 observed, the diameter increase being faster in the juvenile stage. M 362 363 Remarks: Ooster (1860) based the description of this species on two different D 364 specimens (Ooster, 1860: pl. 60, fig. 4, 5). A thorough review of his collection has TE 365 revealed that unfortunately, the larger specimen (Ooster, 1860: pl. 60, fig. 5) is 366 nowadays lost. Nevertheless, and as discussed below, this specimen is reckoned to EP 367 belong to a different, more common species. C 368 AC 369 Ooster (1860) implied that ‘Baculites’ renevieri had been collected in Valanginian strata 370 from Switzerland, together with Bochianites neocomiensis (d’Orbigny, 1842). Were this 371 information correct, Ooster’s species would possess a much earlier stratigraphic range 372 than our specimens. However, a close examination of the drawings of the two syntypes 373 in Ooster’s original work, where they were never explicitly referred to be part of a same 374 individual, reveals that they may belong to two different species. The largest part ACCEPTED MANUSCRIPT 375 (Ooster, 1960: pl. 60, fig. 5) appears to be a fragment of some Anahamulinidae, which 376 was also pointed out by Breistroffer (1952), who ventured an early Barremian age for 377 Ooster’s material. Specifically, the aforementioned form is particularly close to 378 Amorina pictetiformis (Busnardo in Busnardo et al., 2003) (=Anahamulina jourdani 379 sensu Company et al., 2005), typical from the latest Hauterivian (Busnardo et al., 2003). PT 380 Conversely, the smaller fragment (Ooster, 1960: pl. 60, fig. 4) is clearly conspecific RI 381 with the Spanish material shown in the present work. If the two specimens illustrated by 382 Ooster were from the same age, as he implied, and the largest one were indeed part of SC 383 an A. pictetiformis, the stratigraphic range of G. renevieri would be updated to the latest 384 Hauterivian, coinciding with our specimens. 385 U AN 386 Klinger & Kennedy (1979) recorded Bochianites cf. renevieri from the Valanginian of M 387 South Africa with a small fragment bearing a particularly marked constriction (Klinger 388 & Kennedy, 1979: fig. 3, E). Considering the fragmentary nature of their material, and D 389 following the arguments above, we think that the depicted specimen could belong to TE 390 some other species of Valanginian Bochianites, since the constriction follows a slightly 391 flexuous shape, whereas the specimens of G. renevieri studied herein bear straight EP 392 constrictions. C 393 AC 394 Vašíček & Faupl (1999) illustrated a particular specimen (Vašíček & Faupl, 1999: pl. 6, 395 fig. 1) from the upper Hauterivian (‘Angulicostata-Zone’ ≈ Ohmi Zone as used herein) 396 of Austria, which is clearly hamuliconic and possesses an ornamentation pattern on the 397 phragmocone that very much resembles both our material and Ooster’s smaller syntype. 398 They doubtfully determined it as ?Anahamulina sp. Due to ornamental and stratigraphic 399 coincidence, it could be regarded to belong to the same species as the material studied in ACCEPTED MANUSCRIPT 400 the present work. However, at the end of the proversum some thin ribs can be 401 recognised. Therefore, the specimen may indeed correspond to some Anahamulinidae, 402 the particular phragmocone ‘ornamentation’ being probably caused by the dissolution of 403 the material surrounding the chambers (Z. Vašíček, pers. comm., 2019). 404 PT 405 Avram (2001) identified a small fragment (Avram, 2001: pl. 1, fig. 3) from the early RI 406 Hauterivian of Romania as Bochianites cf. renevieri. The poor preservation of that 407 specimen makes it difficult to emit any decisive conclusions on its specific SC 408 determination. 409 410 U Tajika et al. (2017) figured a fragment from Switzerland, collected ex situ (K. Tschanz, AN 411 pers. comm., 2018), with an ornamentation pattern consisting in relatively sharp M 412 constrictions and large interspaces, identified as Bochianitinae indet. (Tajika et al., 413 2017: fig. 9-AH). Because most of the phosphatic ammonites were collected in a D 414 phosphatic conglomerate of lower Barremian (Pulchella–Compressissima Zones) age, TE 415 such time interval was considered for the Swiss specimen. That specimen is herein 416 thought to correspond to G. renevieri, as both size and ornamentation are consistent. It EP 417 illustrates the high ornamental variability in terms of separation of the constrictions (cf. Fig. 2J herein, which shows a specimen with very narrow interspaces at a similar C 418 AC 419 diameter). The only concern is that Tajika et al.’s specimen is notably younger than all 420 the Spanish well-dated material. Two possible hypotheses are put forward to explain 421 this case: [1] Should the specimen indeed belong to the Barremian beds, it would 422 indicate the survival of the species beyond the P. mortilleti Subzone, which is the 423 youngest age in which Spanish G. renevieri have been collected. [2] Tajika et al.’s 424 specimen could in fact belong to another, basal phosphatic and glauconitic bed present ACCEPTED MANUSCRIPT 425 at the locality (Mortilleti Subzone). The aforementioned authors considered it to belong 426 to the lower Barremian because the matrix was seemingly equivalent to the one of those 427 beds. However, re-examination of the matrix of Tajika et al.’s specimen, characterised 428 by a brownish colour and a high quantity of glauconite grains, reveals a strong contrast 429 with the black colour of the phosphatic Barremian specimens. This, added to recent PT 430 findings of phosphatic specimens of Pseudothurmannia cf. pseudomalbosi from the RI 431 nearby Tierwis section (PK.05.A.02.07, Coll. P. Kürsteiner) could indicate that the 432 Tajika et al.’s specimen originally belonged to an upper Hauterivian phosphatic bed SC 433 instead, just like the rest of well-dated specimens studied in the present paper. In any 434 case, the present work does not consider Tajika et al.’s specimen to frame the 435 U stratigraphic range of the species, in hopes that future collections will confirm or deny AN 436 the presence of G. renevieri in the early Barremian. M 437 438 Among the specimens illustrated in this work, almost all the juvenile shafts, similar to D 439 Ooster’s type, have been found in close association with Balearites binelli (Astier, TE 440 1851), Balearites krenkeli (Sarkar, 1955) and Balearites angulicostatus (d’Orbigny, 441 1842), undoubtedly marking their respective subzones, although in different deposits. EP 442 The only supposed subadult fragment (Fig. 2J) has also been collected in the Krenkeli Subzone, in association with its index species. Finally, one fragmentary specimen has C 443 AC 444 been found in association with Pseudothurmannia pseudomalbosi Sarasin & 445 Schöndelmayer, 1901, thus marking the M.ortilleti Subzone. 446 447 448 Glaucoceras aff. renevieri (Ooster, 1860) 449 (Fig. 2L–M) ACCEPTED MANUSCRIPT 450 451 Studied material: Two fragmentary specimens; one from the B. krenkeli Subzone from 452 Can Negret (MBCN 23393) and one from the B. angulicostatus Subzone of Arroyo 453 Gilico (X.V1(–1)56). 454 PT 455 Stratigraphic and geographic distribution: Late Hauterivian, hitherto only found in RI 456 the B. krenkeli and B. angulicostatus Subzones (B. balearis Zone). So far, only known 457 from Spain (the present work). SC 458 459 Diagnosis: Small–?medium size. Heteromorphic shell shape, most probably 460 U baculiconic, although a hamuliconic or ptychoconic shape cannot be entirely ruled out. AN 461 Subcircular section; flanks, dorsum and venter convex. Very simple ornamentation M 462 consisting in shallow and sharp, neatly radial, straight constrictions, with progressively 463 larger interspaces. They apparently cross both the venter and the dorsum, weakening D 464 slightly on the latter. Suture line unknown. TE 465 466 Ontogenetic and intraspecific variation: Even with a small number of specimens, EP 467 some characters can be regarded as variable, such as the separation of the constrictions, which seems to decrease along ontogeny and between specimens, and the depth and C 468 AC 469 sharpness of constrictions, which also seems to be quite variable (see Tab. 2). 470 471 Remarks: G. aff. renevieri differs from the typical form of G. renevieri because the 472 latter bears clearly prosriradiate constrictions, which are also more spaced from each 473 other. Although it could be interpreted as a different species, the authors of the present ACCEPTED MANUSCRIPT 474 paper consider that such a conclusion would be highly presumptuous with a pool of just 475 two fragmentary specimens to compare. 476 477 478 PT 479 5. Discussion RI 480 481 The origination of heteromorphic ammonite lineages, especially those of small size, has SC 482 long been linked to stressful events, such as changes in sea level, trophic or oxygenation 483 conditions. Some examples from the literature include: Ernst et al. (1983) showed that 484 U the late Turonian Hyphantoceras Hyatt, 1900 event corresponds to a transgressive AN 485 interval. Hoedemaeker (1995) claimed that some genera of small heteromorphic M 486 ammonites (Hamulinites Paquier, 1900, Karsteniceras Royo y Gómez, 1945, 487 Protancyloceras Spath, 1924, Leptoceras Uhlig, 1883) were opportunistic and D 488 flourished right after extinction events. Company et al. (1995) also recognised this TE 489 pattern in the genera Hamulinites Paquier, 1900, Tzankoviceras Manolov, 1962, 490 Protancyloceras Spath, 1924 and Leptoceras Uhlig, 1883. Cecca (1997) proposed that EP 491 the cause of the diversification of Protancyloceratina in the Tithonian could be related to an increase of primary productivity, and that the diversification of ‘basal ancyloceratid’- C 492 AC 493 like morphologies near the Valanginian–Hauterivian boundary coincided with an anoxic 494 event, with an increase of food resources in the shallowest parts of the water column 495 (also studied by Reboulet, 2008 in the particular case of Himantoceras Thieuloy, 1965). 496 Lukeneder (2003, 2005) described mass occurrences of Karsteniceras Royo y Gómez, 497 1945 in the Barremian, linked to dysoxic waters. Reboulet et al. (2003) demonstrated 498 that the higher abundance of Valanginian Bochianites Lory, 1898 occurred during ACCEPTED MANUSCRIPT 499 anoxic events, pointing to their opportunistic mode of life. Reboulet et al. (2005) 500 studied the latest Albian Breistroffer interval (OAE1d), and concluded that 501 heteromorphic ammonites diversified themselves and became more abundant during 502 this transgressive, anoxic event. Yet, Guex (2001, 2006) discussed a seemingly opposite 503 case: the origination of some heteromorphic ammonite lineages in response to major PT 504 regressive events related to a reinitialisation of evolutionary clocks and evolutionary RI 505 processes such as proteromorphosis. Additionally, Bert & Bersac (2013) reported that, 506 although the late Barremian Gassendiceras Bert et al., 2006 species acquired SC 507 heteromorphic coilings during a transgressive episode, their most uncoiled morphotypes 508 appeared during an arid interval, which can be related to a diminished fertilisation of the 509 U ocean waters. Therefore, whereas most of the heteromorphic ammonite lineages seem to AN 510 have originated during transgressive events with nutrient-rich waters, there are M 511 particular cases of other lineages becoming heteromorphic during regressive episodes or 512 intervals of low nutrient runoff to the oceans. D 513 TE 514 For the uppermost Hauterivian, Company et al. (2005) located the sequence boundaries 515 of Hardenbol et al. (1998) in a high-resolution biostratigraphic framework. Specifically, EP 516 they placed transgressive maximums at the upper part of the Binelli Subzone and at the Ohmi–Mortilleti Subzones boundary, and regressive maximums at the upper part of the C 517 AC 518 Krenkeli Subzone (SBHa6) and at the upper part of the Picteti Subzone (SBHa7). 519 520 The origination of the genus Glaucoceras seems to be linked to a particular 521 diversification event of heteromorphic ammonites, located at the late B. binelli–early B. 522 krenkeli Subzones (Fig. 3A–B), which coincides with the origination of the genera 523 Monodites Bert, 2009, Paracostidiscus Busnardo (in Busnardo et al., 2003), ACCEPTED MANUSCRIPT 524 Garroniceras Vermeulen et al. 2012a, Amorina Vermeulen, 2005 and Hamulinites 525 Paquier, 1900 (see Supplementary Material 2 for specific references on the species). 526 This event probably corresponds to the transgressive maximum of the late B. binelli 527 Subzone pinpointed by Company et al. (2005), and therefore this case seems consistent 528 with the bulk of examples in the literature: heteromorphic ammonite lineages that PT 529 appear or radiate during transgressive intervals (they have often been argued to be RI 530 opportunistic in nature, see above for references). 531 SC 532 This event can be compared to the more extensively studied Faraoni Ocean Anoxic 533 Event (Company et al., 2005; Föllmi, 2012; Aguado et al., 2014) that occurred at the P. 534 U ohmi–P.mortilleti Subzones boundary, and corresponds to the next and more important AN 535 maximum flooding surface. By then, several small to medium-sized heteromorphic M 536 genera such as Leptoceratoides Royo y Gómez, 1945, Sabaudiella Busnardo (in 537 Busnardo et al., 2003), Bastelia Vermeulen et al., 2012a and Anahamulina Hyatt, 1900 D 538 made their first appearance (Fig. 3A–B), along with the turnover of many other TE 539 ammonoid taxa (Company et al., 2005; Supplementary Material 2 for references on 540 heteromorphic species). That transgressive maximum was accompanied by more humid EP 541 conditions, which increased the nutrient runoff from the continent (Aguado et al., 2014). This phenomenon increased the eutrophication of the ocean waters, leading to C 542 AC 543 the decrease of dissolved oxygen levels, which affected the marine ecosystems. 544 545 546 6. Conclusions 547 ACCEPTED MANUSCRIPT 548 Detailed sampling of different outcrops both from Mallorca and the Betic Cordillera 549 allowed the recovery of fragments of 13 ammonoid fossil shells attributed to the newly 550 created genus Glaucoceras gen. nov. The bed-by-bed collecting process (in the case of 551 Arroyo Gilico) or the direct association with the respective index fossils (in the 552 Mallorcan deposits) allowed to provide precise biostratigraphic range for each sample. PT 553 Two morphotypes can be distinguished within this new genus, which appears to be RI 554 quite rare, since only two specimens had been illustrated by previous authors. 555 Glaucoceras renevieri, a new combination for Ooster (1860)’s species Baculites SC 556 Renevieri, seems widespread from the B. binelli Subzone to the P. mortilleti Subzone. 557 Contrarily, Glaucoceras aff. renevieri appears to span from the B. krenkeli Subzone to 558 U the B. angulicostatus Subzone. This data improves the knowledge of the heteromorphic AN 559 ammonoid fauna during the late Hauterivian, which is a period of great diversification M 560 of these groups. Specifically, the bulk of heteromorphic genera were originated in two 561 specific events, coinciding with the consecutive transgressive maximums in the late B. D 562 binelli–early B. krenkeli Subzones and in the P. ohmi–P. mortilleti Subzones boundary TE 563 (Faraoni Ocean Anoxic Level). This allows to put forward the hypothesis that latest 564 Hauterivian heteromorphic ammonoids became more diverse during eustatic EP 565 maximums, when ocean waters flooded the continental shelves, resulting in the increase of nutrient runoff to the sea, favouring the development of epipelagic forms. However, C 566 AC 567 further research will be required to ascertain the specific traits that made several 568 independent lineages of heteromorphic ammonites ecologically more successful in 569 nutrient-rich waters, to determine if there exist differences in evolutionary patterns 570 between small- and large-sized heteromorphic ammonoids, and to prove whether both 571 size classes were opportunistic or not (hitherto, only small-sized heteromorphs were 572 regarded as such). ACCEPTED MANUSCRIPT 573 574 575 7. Acknowledgements 576 577 The authors of this work express their gratitude to Dr. Miguel Company (Universidad PT 578 de Granada, Spain) for allowing us to check their collections and use and illustrate their RI 579 specimens in this paper, as well as for his thoughtful insight on several parts of this 580 manuscript. To Dr. Didier Bert (Réserve naturelle nationale géologique de Haute- SC 581 Provence, Université de Rennes-1 and Laboratoire du Groupe de recherche en 582 Paléobiologie et biostratigraphie des Ammonites, France) and Dr. Ottilia Szives 583 U (Hungarian Natural History Museum, Hungary), reviwers of this paper, for their most AN 584 useful comments, that contributed to improving the original text. To Dr. Zdeněk M 585 Vašíček (Institute of Geonics, Academy of Sciences of the Czech Republic, Czech 586 Republic), for his clarifying remarks on a particular specimen figured in one of his D 587 papers that was considered herein. To Karl Tschanz for information on Tajika et al. TE 588 (2017)’s Bochianitinae indet. specimen. To Peter Kürsteiner (Naturmuseum St. Gallen, 589 Switzerland) for the access to his phosphatic specimen of Pseudothurmannia cf. EP 590 pseudomalbosi. To Pere Miralles, for the help in confection of one of the maps. We also thank Sebastià Matamalas for his aid in the fieldwork and collection of some of the C 591 AC 592 specimens from Mallorca studied herein. This research did not receive any specific 593 grant from funding agencies in the public, commercial, or not-for-profit sectors. 594 595 596 8. References 597 ACCEPTED MANUSCRIPT 598 Agassiz, L. (1847). Introduction to the study of Natural History, in a series of lectures 599 delivered in the Hall of the College of Physicians and Surgeons, New York. Greley & 600 McElrath. 58 pp. 601 602 Aguado, R.; Company, M.; O’Dogherty, L.; Paloma, I.; Sandoval, J. & Tavera, J.M. PT 603 (2008). Integrated stratigraphy of the uppermost Hauterivian–Lower Barremian pelagic RI 604 sequence of Arroyo Gilico (Betic Cordillera, SE Spain). Berichte der Geologischen 605 Bundesanstalt, 74: 36–37. SC 606 607 Aguado, R.; Company, M.; O’Dogherty, L.; Sandoval, J. & Tavera, J.M. (2014). 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Denkschriften 896 der Kaiserlichen Akademie der Wissenschaften, Wien, Mathematisch- 897 naturwissenschaftliche Classe, 46: 127–290 (1–166) + 32 pls. 898 PT 899 Vašíček, Z. (1972). Ammonoidea of the Těšín–Hradiště Formation (Lower Cretaceous) RI 900 in the Moravskoslezské Beskydy Mts. Rozpravy Ústředního Ústavu Geologického, 38: 901 104 pp. + 16 pls. SC 902 903 Vašíček, Z. (1999). Comments on some new occurrences of heteromorph ammonites in 904 U the Lower Cretaceous (Late Valanginian – Early Barremian) of the Eastern Alpine and AN 905 Western Carpathian systems. Scripta Geologica, special issue 3: 215–227. M 906 907 Vašíček, Z. & Faupl, P. (1999). Zur Biostratigraphie des Schrambachschichten in der D 908 Reichraminger Decke (Unterkreide, oberösterreichische Kalkalpen). Abhandlungen der TE 909 Geologischen Bundesanstalt, 56(2): 593–624. 910 EP 911 Vašíček, Z. & Hoedemaeker, P.J. (2003). Small Berriasian, lower Valanginian and Barremian heteromorphic ammonites from the Río Argos succession (Caravaca, C 912 AC 913 southeast Spain). Scripta Geologica, 125: 11–33. 914 915 Vašíček, Z. & Wiedmann, J. (1994). The Leptoceratoidinae: small heteromorph 916 ammonites from the Barremian. Palaeontology, 37(1): 203–239. 917 918 Vera, J.A. (2004) (ed.). Geología de España. IGME, Madrid. 883 pp. + 2 maps. ACCEPTED MANUSCRIPT 919 920 Vermeulen, J. (2005). Boundaries, ammonite fauna and main subdivisions of the 921 stratotype of the Barremian. In: Adatte, T.; Arnaud-Vanneau, A.; Arnaud, H.; Blanc- 922 Aletru, M.-C.; Bodin, S.; Carrio-Schaffauser, E.; Föllmi, K.; Godet, A.; Raddadi, M.C. 923 & Vermeulen, J. (eds.). The Hauterivian lower Aptian sequence stratigraphy from Jura PT 924 platform to Vocontian basin: a multidisciplinary approach. Field-trip of the 7th RI 925 International symposium on the Cretaceous (September 1–4, 2005). Géologie Alpine, 926 Colloques et Excursions, 7: 147–173. SC 927 928 Vermeulen, J. (2006). Nouvelle classification à fondement phylogénétique des 929 U ammonites hétéromorphes du Crétacé inférieur. Annales du Muséum d’Histoire AN 930 Naturelle de Nice, 21: 137–178. M 931 932 Vermeulen, J. (2009). Taxa noveaux ou peu connus de la sous-famille des D 933 Anahamulininae (Turritilina, Ptychoceratoidea, Hamulinidae). Annales du Muséum TE 934 d’Histoire Naturelle de Nice, 24(2): 103–131. 935 EP 936 Vermeulen, J. (2010a). Nouvelle tripartition de la famille des Anahamulinidae Breistroffer, 1952 (Turrilitina, Ptychoceratoidea) et description de quelques taxa du C 937 AC 938 Barrémien inférieur du sud-est de la France. Annales du Muséum d’Histoire Naturelle 939 de Nice, 25: 61–91. 940 941 Vermeulen, J. (2010b). Sur deux nouvelles espèces de la famille des Anahamulinidae 942 Breistroffer, 1952 (Turrilitina, Ptychoceratoidea). Riviera Scientifique, 94: 107–116. 943 ACCEPTED MANUSCRIPT 944 Vermeulen, J. & Vašíček, Z. (2011). Espèces d’ammonites nouvelles ou peu connues de 945 la famille des Anahamulinidae Breistroffer, 1952 (Turrilitina, Ptychoceratoidea). 946 Annales du Muséum d’Histoire Naturelle de Nice, 26: 47–94. 947 948 Vermeulen, J.; Cecca, F. & Kruta, I. (2007). Nouveaux taxa de la famille des PT 949 Hamulinidae Gill, 1871 (Ammonitida, Turrilitina, Ptychoceratoidea). Annales du RI 950 Muséum d’Histoire Naturelle de Nice, 22: 1–25. 951 SC 952 Vermeulen, J.; Lazarin, P.; Lépinay, P.; Leroy, L. & Mascarelli, E. (2010a). Nouvelles 953 données nomenclaturales sur les Ptychoceratoidea (p. parte) du sud-est de la France et 954 U description de quelques taxa du Barrémien supérieur de la famille des Anahamulinidae. AN 955 Annales du Muséum d’Histoire Naturelle de Nice, 25: 19–59. M 956 957 Vermeulen, J.; Company, M.; Sandoval, J. & Tavera, J.M. (2010b). Sur deux espèces D 958 nouvelles d’ammonites de l’Hauterivien supérieur du sud-est de l’Espagne. Annales du TE 959 Muséum d’Histoire Naturelle de Nice, 25: 93–104. 960 EP 961 Vermeulen, J.; Lazarin, P.; Lépinay, P.; Leroy, L.; Mascarelli, E.; Meister, C. & Menkveld-Gfeller, U. (2012a). Ammonites (Ancyloceratina, Turrilitina) nouvelles ou C 962 AC 963 peu connues de l’Hauterivien supérieur. Annales du Muséum d’Histoire Naturelle de 964 Nice, 27: 271–318. 965 966 Vermeulen, J.; Lazarin, P.; Lépinay, P.; Leroy, L. & Mascarelli, E. (2012b). Nouvelles 967 données sur l’évolution et la classification de quelques Anahamulinidae Breistroffer, 968 1952 (Turritilina, Ptychoceratoidea). Riviera Scientifique, 96: 79–95. ACCEPTED MANUSCRIPT 969 970 Vermeulen, J.; Lazarin, P.; Lépinay, P.; Leroy, L. & Mascarelli, E. (2013a). Taxa 971 d’ammonites nouveaux ou peu connus de l’Hauterivien et du Barrémien du sud-est de la 972 France. STRATA, 2nd series: mémoires, 48. 58 pp. 973 PT 974 Vermeulen, J.; Lazarin, P.; Lépinay, P.; Leroy, L. & Mascarelli, E. (2013b). Le genre RI 975 Defayella gen. nov. et ses implications taxinomiques (Turrilitina, Ptychoceratoidea, 976 Anahamulinidae). Riviera Scientifique, 97: 83–96. SC 977 978 Vermeulen, J.; Lukeneder, A.; Meister, C. & Vašíček, Z. (2014a). New data on four 979 U species of ammonites created by V. Uhlig in 1883 (Ammonitida, Turrilitina, AN 980 Hamulinoidea). Annales du Muséum d’Histoire Naturelle de Nice, 29: 1–25. M 981 982 Vermeulen, J.; Lazarin, P.; Lépinay, P.; Leroy, L. & Mascarelli, E. (2014b). Ammonites D 983 du Barremien du Sud-Est de la France (Ammonitina, Ancyloceratina, Turritilina). TE 984 STRATA, 2nd series: mémoires, 50. 95 pp. 985 EP 986 Weidmann, M.; Homewood, P.; Morel, R.; Berchten, J.-D.; Bucher, H.; Burri, M.; Cornioley, J.-R.; Escher, P.; Rück, Ph.; Tabotta, A. & Zahner, Ph. (1993). Atlas C 987 AC 988 géologique de la Suisse au 1:25ʼ000, feuille 1244 Châtel-St-Denis. Notice explicative 989 par M. Weidmann. Service hydrologique et géologique national, Berne: 55 pp. 990 991 Wiedmann, J. (1962a). Unterkreide-Ammoniten von Mallorca 1. Lieferung: 992 Lytoceratina, Aptychi. Akademie der Wissenschaften und der Literatur. Abhandlungen 993 der Mathematisch-Naturwissenschaftlichen Klasse, 1: 1–148 + 10 pls. ACCEPTED MANUSCRIPT 994 995 Wiedmann, J. (1962b). Ammonites du Crétacé inférieur de Majorque (Baléares). 1. 996 Partie: Lytoceratina et Aptychi. Boletín de la Sociedad de Historia Natural de Baleares, 997 8: 3–54 + 10 pls. 998 PT 999 Wiedmann, J. (1964). Unterkreide-Ammoniten von Mallorca. 2. Lieferung: RI 1000 Phylloceratina. Akademie der Wissenschaften und der Literatur. Abhandlungen der 1001 Mathematisch-Naturwissenschaftlichen Klasse, 4 (1963): 151–264 + 11 pls. SC 1002 1003 Wiedmann, J. (1966). Stammesgeschichte und System der posttriadischen 1004 U Ammonoideen, etn Überblick (2. Teil). Neues Jahrbuch für Geologie und AN 1005 Palaeontologie, Abhandlungen, 127: 13–81 + 4 pls. + 34 figs. M 1006 1007 Wiedmann, J. (1967). Ammonites du Crétacé inférieur de Majorque (Baléares). 2.ª D 1008 Partie: Phylloceratina. Boletín de la Sociedad de Historia Natural de Baleares, 13: 3–40 TE 1009 + 11 pls. C EP AC 1 Table 1: Stratigraphic framework used herein (from Matamales-Andreu & Company, 2018) and distribution of the studied species ACCEPTED MANUSCRIPT 2 in each of the considered localities. Veveyse de Châtel not included because the stratigraphic position of the only specimen is 3 uncertain. Dark blue (×): Gl. renevieri; Orange (+): Gl. aff. renevieri. Age Locality Stage Zone Subzone Arroyo Gilico Can Negret Biniamar Son Macià Manacor P. picteti Pseudothurmannia P. mortilleti × ohmi Late P. ohmi Hauterivian B. angulicostatus × + pro parte Balearites B. krenkeli × + × × PT balearis B. binelli × B. balearis 4 RI 5 6 Table 2: Biometric measurements of all the studied specimens. Width and length taken in cm. SC Minimum– Maximum Number of Constrictions Specimen Species Growth rate maximum width length constrictions per cm NMBE 5007731 Gl. renevieri 0.24–0.58 4.73 0.072 14 3 U MBCN 23389 Gl. renevieri 0.31–0.34 1.80 0.017 11 6 MBCN 23392 Gl. renevieri 0.18–0.38 2.04 0.098 18 8 AN MBCN 23390 Gl. renevieri 0.51–1.00 5.97 0.082 24 4 MBCN 23391 Gl. renevieri 0.35–0.56 2.41 0.087 9 3 MBCN 23395 Gl. renevieri 0.27–0.32 1.22 0.041 10 8 MBCN 23394 Gl. renevieri 0.33–0.34 1.18 0.008 7 5 M X.V1 (–16) 9 Gl. renevieri 0.15–0.30 1.75 0.086 24 13 X.V1 (–10) 21 Gl. renevieri 0.20–0.24 1.04 0.038 17 16 X.V1 (–2) 57 Gl. renevieri 0.25–0.30 0.99 0.051 7 7 X.V1 (–1) 17 Gl. renevieri 0.29–0.32 1.33 0.024 16 12 D MBCN 23393 Gl. aff. renevieri 0.42–0.52 1.15 0.087 9 7 X.V1 (–1) 56 Gl. aff. renevieri 0.29–0.50 1.53 0.137 19 12 TE 7 8 EP 9 Figure 1. Location of the studied sites. A: Geological map of SW Switzerland with the location of Veveyse de Châtel (a). B: 10 Geological map of Mallorca with the four location of Biniamar (b), Can Negret (c), Manacor (d) and Son Macià (e). C: Studied C 11 section in the locality of Biniamar. D: Geological map of the Betic Cordillera with the location of Arroyo Gilico (f). AC 12 13 Figure 2. A–K: Glaucoceras renevieri (Ooster, 1860). A: Specimen NMBE 5007731, Holotype, upper Hauterivian, Veveyse de 14 Châtel. B: Sample MBCN 23391 in association with B. “majoricensis”, Krenkeli Subzone, Manacor. C: Sample MBCN 23392 in 15 association with Phyllopachyceras winkleri, Krenkeli Subzone, Biniamar. D: Sample MBCN 23394 in association with B. 16 “majoricensis”, Krenkeli Subzone, Biniamar. E: Specimen MBCN 23395, Krenkeli Subzone, Biniamar. F: Specimen X.V1 (–1) 17 17, Angulicostatus Subzone, Arroyo Gilico. G: Specimen X.V1 (–16) 9, Binelli Subzone, Arroyo Gilico. H: Specimen X.V1 (– 18 10) 21, Krenkeli Subzone, Arroyo Gilico. I: Sample MBCN 23389 in association with P. pseudomalbosi, Mortilleti Subzone, Son 19 Macià. J: Specimen MBCN 23390, Krenkeli Subzone, Manacor. K: Specimen X.V1 (–2) 57, Angulicostatus Subzone, Arroyo 20 Gilico. L–M: Glaucoceras aff. renevieri. L: Specimen X.V1 (–1) 56, Angulicostatus Subzone, Arroyo Gilico. M: Specimen ACCEPTED MANUSCRIPT 21 MBCN 23393, Krenkeli Subzone, Can Negret. Scale bar: 10 mm. 22 23 Figure 3. A: Stratigraphic distribution of heteromorphic ammonoid genera for the latest Hauterivian (B. balearis and P. ohmi 24 Zones). Data on sources of information on stratigraphic ranges can be consulted in Supplementary Material 2. Biozonation 25 framework as in Tab. 1 herein. Sea level information taken from Company et al. (2005). Dashed lines indicate possible phyletic PT 26 relationships. Question marks represent unknown parts of the ammonites. The star marks the possible diversification event of 27 heteromorphic ammonites at the B. binelli–B. krenkeli Subzones boundary. B: Stacked area chart showing the changes of diversity RI 28 of heteromorph ammonoids over the latest Hauterivian, using the data presented in the Supplementary Material 2. Arrows indicate 29 the two diversification events. U SC AN M D TE C EP AC ACCEPTED MANUSCRIPT PT RI U SC AN M D TE EP C AC ACCEPTED MANUSCRIPT PT RI U SC AN M D TE EP C AC ACCEPTED MANUSCRIPT PT RI U SC AN M D TE EP C AC ACCEPTED MANUSCRIPT • The genus Glaucoceras gen. nov. is described to include the poorly-known species Baculites Renevieri Ooster, 1860. Glaucoceras renevieri is revised and compared with other similar taxa. • The stratigraphic range of Glaucoceras renevieri is updated, revealing that it appears between the Balearites binelli Subzone (Balearites balearis Zone) and the Pseudothurmannia mortilleti Subzone (Pseudothurmannia ohmi Zone) of the latest PT Hauterivian (Early Cretaceous). RI • The palaeoenvironmental event that is thought to have led to the origination of this genus is further explored, concluding that it corresponded to a transgressive maximum SC that occurred at the Balearites binelli–Balearites krenkeli Subzones (Balearites balearis Zone). U • Two successive transgressive events in the latest Hauterivian are interpreted as the AN triggers of the diversification of many other heteromorphic ammonoid lineages. M D TE C EP AC