The Phyletic Origin of Zigzagiceratinae
NEW SPECIES OF FRANCHIA AND PROTOZIGZAGICERAS (AMMONOIDEA, MIDDLE JURASSIC): THE PHYLETIC ORIGIN OF ZIGZAGICERATINAE
by SIXTO R. FERNANDEZ-LOPEZ1* and GIULIO PAVIA2
lDepartarnento de Paleontologia, Facultad de Ciencias Geol6gicas, Universidad Complutense de Madrid, Calle Jose Antonio Novais 2, 28040, Madrid, Spain; e-mail: [email protected] 'Dipartimento di Scienze della Terra, via Valperga Caluso 35, 10125, Torino, Italy; e-mail: [email protected]
Abstract: Three genera and seven species belonging to the shell. In contrast, rapid proterogenesis originated and diver subfamily Zigzagiceratinae (family Perisphinctidae) are sified the earliest Bathonian zigzagiceratin lineages, giving described from the Lower Bathonian of France and Saudi paedomorphic members, commonly neotenic and more scar Arabia. Intraspecific dimorphism is recognized. A revision of cely progenetic. Procerites-Siemiradzkia seems to be the old the genus Franchia proposed by Sturani (1967), based on the est zigzagiceratin member in the French Subalpine, Iberian syntypes and new specimens from south-east France, is and Lusitanian basins, branched off by paedomorphosis from presented. Franchia arkelli Sturani, Franchia subalpina sp. leptosphinctins at the Bajocian-Bathonian transition. The nov., Protozigzagiceras torrensi (Sturani), Protozigzagiceras Mediterranean-Caucasian genera Franchia, Zigzagiceras, Zig tethycum sp. nov., Protozigzagiceras flexum sp. novo and Pro zagites and Wagnericeras branched from successive species of tozigzagiceras densum sp. novo are described from the Digne Protozigzagiceras, in turn, a direct derivative of Procerites. Castellane region of south-east France. Megazigzagiceras The oldest lineages of the clade Zigzagiceratinae evolved by subarabicum, gen. et sp. novo is described from the Dharma iterative, rapid, paedomorphic changes and additional, last region of Saudi Arabia. The successive Early Bathonian spe ing, peramorphic modifications during the Early Bathonian. cies of Franchia and Protozigzagiceras herein identified in West Tethyan areas, as members of the Mediterranean-Cau Key words: Ammonites, Perisphinctidae, Western Tethys, casian Subrealm, formed lasting separate peramorphoclines Ethiopian Province, Bathonian Global Stratotype Section and characterized by increasing hydrodynamic coiling of the Point.
THE Early Bathonian genera Franchia and Protozigzagiceras determined by Sturani (1967) as Franchia arkelli and Zig characterize the Zigzag Zone, in particular the Parvum and zagiceras torrensi, assuming a high variability of sutural Macrescens subzones of the French Subalpine Basin complexity in Franchia and of zigzag development in Zig (Fig. lA), although they are scarce components of the zagiceras. Sturani (1967) and Torrens (1987) established the ammonite fossil assemblages «5.0 per cent). The accurate biochronostratigraphical foundations for the Lower Batho biochronostratigraphy of these Perisphinctidae is crucial nian ammonoid succession of Digne-Castellane region. for calibration of the basal Bathonian Zigzag Zone and cor Complementary biostratigraphical information on Zigzagi relation of the Bathonian Global Stratotype Section and ceratinae was developed in graduate theses by Innocenti Point (GSSP), currently defined in the Ravin du m�s Section, (1975), Puma (1975) and Romeo (1999). Innocenti et al. Bas Auran area (Fig. lB, Fernandez-Lopez et al. 2009b). (1990) proposed the boundary stratotype of the Bathonian The specimens belonging to the subfamily Zigzagicerati Stage based on a Bas Auran section. Fernandez-Lopez nae studied herein have been collected, bed-by-bed, in sev (2007) presented a taphonomic analysis of the ammonoid eral sections from the 'Marno-calcaires a Cancellophycus' succession on the Bas Auran area at the Bajocian-Bathonian Formation in south-east France, during the last five dec boundary and the palaeoenvironmental interpretation in ades. Several tens of in situ Lower Bathonian Franchia and terms of sequence stratigraphy. Fernandez-Lopez et al. (2007) Protozigzagiceras from Digne-Castellane region have been published new data about the youngest members of the identified to species level. Most of these specimens were Bigotitinae and the oldest members of the Zigzagiceratinae, Panboreal Realm • Paris
* Mediterranean Caucasian Subrealm
*
East Indo-Pacific Subrealm Pacific Ravin des Robins Subrealm " sect o �I l :avin d'Auran Section Tethys Les B/aches I ' Panthalassa La B/ache Section Realm 5 km 60 19' E N - -- Castellane * French Subalpine Basin * Arabian Platform A
FIG. 1. Geographical positions of the localities mentioned in text. A, palaeogeographical position of the French Subalpine Basin in relation to the Arabian Platform during the Early Bathonian; palaeogeography modified from Fernandez-Lopez et al. (2009b), and Gol onka (2011); palaeobiogeographical units according to Westermann (2000b). B, location map of the stratigraphical sections in the Digne-Castellane region (SE France). of biochronostratigraphical importance for the subdivision Paleontologia, Universita di Torino (Italy), currently and correlation of the basal Bathonian Zigzag Zone. Pavia stored in the Dipartimento di Scienze della Terra. Speci et al. (2008) revised new information on the ammonoid mens are marked with the acronym PU and a sequential biozonation in the Bas Auran area. Fernandez-Lopez et al. registration number. Source of sampling, section and bed (2009a) formally proposed the Rabin du Bes Section as the number indicated by abbreviations: BA, sections of the GSSP of the Bathonian, and synthesized the regional bio Bas Auran area (lithostratigraphical column of Sturani chronostratigraphical scales available for the Bajocian 1967); RB, Ravin du Bes Section; RA, Ravin d'Auran Sec Bathonian boundary. The Bathonian GSSP in the Ravin du tion; RR, Ravin des Robines. The specimen from Dharma, Bes Section was approved by the International Commis Saudi Arabia, of Figure 19, is housed at the Claude sion of Stratigraphy (ICS) in June 2008 and ratified by the Bernard University of Lyon (France). International Union of Geological Sciences (lUGS) in July 2008 (Fernandez-Lopez et al. 2009b). Ammonite abbreviations and measurements. M, macro The aims of this study were: (1) to provide a revision of conch; m, microconch; D, maximum shell diameter; H, the genus Franchia proposed by Sturani (1967) based on whorl height; HID, ratio of whorl height to diameter; the syntypes and new specimens from south-east France, W, whorl width; W/D, ratio of whorl width to diameter; giving emphasis to the dimorphism of Zigzagiceratinae and U, umbilical diameter; UlD, ratio of umbilicus to diam its possible phyletic relationships with Leptosphinctinae or eter; W/H, ratio of whorl width to whorl height; Ni/2, Bigotitinae; and (2) to interpret the palaeobiogeographical internal ribs per half whorl; Ne/2, external ribs per half patterns and evolutionary trends of the oldest Zigzagicera whorl; I, ratio of external to internal ribs. The measure tinae. The main new contribution concerns the phylogenetic ments presented in Fernandez-Lopez and Pavia (2013) relationships between Early Bathonian lineages of Zigzagi are given in mm and were measured at maximum shell ceratinae and the origination of the clade. diameter.
SPECIMEN REPOSITORY AND SYSTEMATIC PALAEONTOLOGY ABBREVIATIONS This published work and the nomenclatural acts it con Institutional abbreviations. The material described herein tains have been registered in Zoobank: http://zoobank.org/ is housed at the collection of the Museo di Geologia e References/F55 76F2D-3C69-4CED-A6FD-IA4D677814CF Class CEPHALOPODA Cuvier, 1798 classification (Callomon in Donovan et al. 1981; Mangold Order AMMONITIDA Fischer, 1882 1988; Fernandez-Lopez et al. 2007). Superfamily PERISPHINCTOIDEA Steinmann, 1890 The subfamily Zigzagiceratinae is known in the Tethys Family PERISPHINCTIDAE Steinmann, 1890 Panthalassa Realm, mainly from the Bathonian. In Eur ope, it is a common component of Lower Bathonian Remarks. The family Perisphinctidae is characterized by assemblages (Mangold 1997). In North America, members planulate shells, with retracted suspensive-lobe and egres of this subfamily are known from deposits of the Upper sive coiling at maturity, differentiated in two dimorphic Bajocian - Lower Bathonian transition as relatively scarce groups: lappetted microconchs, with short body chamber components of certain ammonite assemblages (Wester ribbed to the end, and simple-aperture macroconchs with mann 1992; Evenchick et al. 2010). Very scarce records smooth or distantly ribbed, long body chamber (near are known from Iran, Saudi Arabia, Chile, Argentina and 360 degrees; Lominadze et al. 1993; Shevyrev 2006; Page Russia (Seyed-Emami et al. 1985, 1989; Poulton et al. 2008; Howarth 2013). 1992; Groschke and von Hillebrandt 1994; Riccardi and Westermann 1999; Beznosov and Mitta 2000; Sey et al.
Subfamily ZIGZAGICERATINAE Buckman, 1920 2004; Enay et al. 2007; Shams and Seyed-Emami 2010). The stratigraphical distribution of the oldest genera of [as Zigzagiceratidae Buckman, 1920, TA-Ill, p. 30] [= Zigzagi Zigzagiceratinae in Bas Auran area (Ravin d' Auran and ceratinae Schindewolf, 1925, p. 319, 1962, p. 520; =Siemiradzkii Ravin du Bt�s sections) is shown in Figure 2. nae Westermann, 1958, p. 75; = Gradlisphinctinae Beznosov, The diverse ontogenetic patterns of lateral ribs and me 1982, p. 54] gastriae in the zigzag-stadium among Early Bathonian Mediterranean-Caucasian zigzagiceratins correspond to Remarks. The subfamily Zigzagiceratinae comprises an ordered multistate character that may be coded in the micro- and macroconchs of small to large size (generally following three categories: 30--600 mm), planorbicones to discocones. The whorls 1. Zigzag-state 1: shortly developed in the nucleus up to increase by segments between more or less marked a few mm in diameter, composed of proradiate to pseudoconstrictions or constrictions. Depressed whorl subradiate, straight to slightly bent ribs of variable section on the inner and middle whorls becomes strength and spacing (e.g. present in Procerites and rounded elliptical and more compressed on the outer Siemiradzkia; Fig. 3; Sturani 1967, pI. 18, fig. 3; Hahn whorls. The subfamily is mainly characterized by pseudo 1969, pI. 4, fig. 4, pI. 9, fig. 4). coronate early whorls that may be followed by a peri 2. Zigzag-state 2: relatively well developed in the inner sphinctoid stage, with blunt, bifurcate or trifurcate and intermediate whorls of the phragmocone (at least costae, lacking smooth band or groove on external in the microconchs), firstly composed of proradiate region. The pseudocoronate stage, so-called zigzag-sta to subradiate, straight to slightly bent ribs of variable dium or zigzag-ornament, may be wmposed of parabolic strength and spacing; then appear proradiate to sub nodes, parabolic ribs, constrictions or pseudoconstric radiate, sigmoid ribs of greater strength upflank and tions, megastriae, sigmoid ribs of greater strength up variably spaced; and finally with occasional proradiate flank, and sharp ribs of variable strength and spacing to subradiate parabolic ribs of variable spacing that (d'Orbigny 1846 in 1842-1851, pI. 129, figs 9-11; Sie pass over the venter radially or with a gentle forward miradzki 1898-1899; Buckman 1920 in 1909-1930, pI. inclination (e.g. in Franchia and Protozigzagiceras; 153; Arkell et al. 1957; Arkell 1958 in 1951-1958, pI. 20- Fernandez-Lopez et al. 2007, figs 10-12). 22; Sturani 1967; Hahn 1969, pI. 2, figs 3-5, pI. 9, fig. 3; 3. Zigzag-state 3: developed in the phragmocone, and Mangold 1971; Torrens 1987). These ornamental features body chamber of the microconchs, firstly composed are due to resorption of the shell or local perturbation of of proradiate to sub radiate sigmoid ribs, of variable the apertural growth field controlling the coiling and fur strength and spacing and of greater strength upflank ther growth of the shell during early-ontogenetic, imma and sharp, parabolic ribs; later, subradiate to rursira ture or preadult growth stages (Bucher et al. 1996; diate, distant and sharp, parabolic ribs, projecting Hammer and Bucher 2005; Doguzhaeva 2012). The evolu sharply forward on ventrolateral shoulder, polyfurcate tionary appearance of a new morphological feature in and bent forward over the venter (e.g. in Zigzagiceras early ontogeny, such as the so-called zigzag-stadium, is a and Procerozigzag; Arkell 1958 in 1951-1958, pI. characteristic of the multiple phyletic lineages of Early 20-22; Hahn 1969, pI. 2, figs 3-5, pI. 9, fig. 3). Bathonian Zigzagiceratinae, distinguishing them from The style of ribbing interpreted by Sturani (I967) as of those of Leptosphinctinae and Bigotitinae. This criterion Zigzagiceras, which passes through a stage transitional to also corroborates the dimorphism of each phyletic lineage Siemiradzkia before the parabolic nodes fade out, corre and allows the development of a primarily phylogenetic sponds to the zigzag-state 2 herein distinguished. Accord- On the basis of the syntypes of Franchia studied by Lower Bathonian Stage Sturani (1967) and new specimens from south-east 'Marno-calcaires a Cancellophycus'Formation Terres Noires' Fm. France, as well as a specimen from Saudi Arabia men Thickness (m) tioned by Enay and Mangold (1994, Enay et al. 1987), in Sturani 1967 RB level seven species and three genera are described and inter Ravin preted below, belonging to Franchia, Protozigzagiceras and du Bes Section Megazigzagiceras. The genera of Zigzagiceratinae men Thickness (m) tioned in text are listed in Table 1.
RA level Ravin d'Auran Section Genus FRANCHIA Sturani, 1967 , c::::J Limestone tie Franchia sp. [M & m] _ Marlstone Type species. Zigzagiceras (Franchia) arkelli Sturani, 1967 (p. 52, tie F. arkel/iSTURANI [M & m] pI. 16, fig. 5, pI. 18, figs 2, 4), by monotypy. F. suba/pina sp. novo [M & m] , :- ��. • • ;Protozigzagicerassp. [m & M] , ' Type level. Lower Bathonian, Zigzag Zone, Macrescens Subzone. ___ P. torrensi (STURANI) [m & M] : Level BAl2 of the 'Marno-calcaires it Cancellophycus' Formation , • P. postpal/ubrum(WETZEL) [m & M] in Bas Auran area (Sturani 1967) and level RA033 of Ravin : .....:. P.tethycum sp. novo [m & M] d'Auran Section (Pavia et al. 2008), indicated in Figure 2. : � P. flexum sp. novo [m & M] � P. densum sp. novo [m] Type locality. Bas Auran. , . Zigzagiceras sp. [m & M] � Wagnericeras sp. [M & m] ...-- ---..� Type region. AIpes de Haute Provence (Bas Auran and Chau , Zigzagites sp. [M & m] • �" 'I don), south-east France. , ' Proceritessp. [M & m] :.. . ---i----.-...... -jo�. Sturani Bh. ' Mondeg. ,Prot! Biohorizon Diagnosis. Conchs of small or medium sIze (40- Parvum Subzone I Macrescens : Recinc.I Ten � Subzone 260 mm). Moderately evolute to moderately involute plan : Aurigerus Zigzag Zone : Zone orbicones. Zigzag-state 2 developed in the phragmocone, Zone ' firstly composed of proradiate to subradiate ribs of vari FIG. 2. Stratigraphical distribution of Lower Bathonian Zigzagi able strength and spacing; then appear proradiate to sub ceratinae (Franchia, Protozigzagiceras, Zigzagiceras, Wagnericeras, radiate, sigmoid ribs of greater strength upflank with Zigzagites and Procerites) in Ravin d'Auran and Ravin du Bes sec variable spacing; and finally, occasional proradiate to sub tions (France) according to the systematic revision presented in radiate, sharp, parabolic ribs of variable spacing. Coarse this paper. The range lines of the genera display the occurrence of fragmentary specimens of undeterminable species and their strati and blunt costae generally bifurcate. Relatively simple graphical constancy through the sections. Logs from Fernandez suture line, with broad lateral lobe. Lopez (2007) and Pavia et al. (2008). Standard chronostrati graphical zonation from Pavia et al. (2008) and Fernandez-Lopez Remarks. Franchia has been regarded as macroconchiate et al. (2009a, b). For location of sites, see Figure lB. with a simple peristome. The counterpart micro conch of Franchia, with lappets, is described here for the first time. Besides the type species, a new species is described and ing to the biochronostratigraphical information obtained interpreted below. Franchia [M & m] is closely homeo in French Subalpine, Iberian and Lusitanian basins morphic with some Lower Bathonian Bigotites Nicolesco (Fig. 2; Fernandez-Lopez 2000; Fernandez-Lopez et al. & (1918; Bigotella petri 2007, 2009a; Pavia et al. 2008), these ontogenic modifica [M m] type species: Nicolesco, 1917, p. 167, pI. 1, figs 4-5), especially in the coiling, the tions of zigzag-states in Early Bathonian zigzagiceratins of segmentary development of the whorls, the prominent pri the western Tethys represent successive, derived or apo mary ribs and the relatively simple suture line, but display morphic conditions (Gould 1977, 2002; Dommergues zigzag-state 2. Other Early Bathonian Mediterranean et al. 1989; Dommergues 1990; Landman et al. 1991; Caucasian macroconchiate zigzagiceratins display more Davis et al. 1996; Guex et al. 2003; Guex 2006). There complex suture lines. The leptosphinctins Planisphinctes fore, these three early ontogenic, zigzag-states identified Buckman [m] (1922; type species: P. planilobus Buckman, in the clade Zigzagiceratinae represent chronostratigraphi 1922 in 1909-1930, TA-IV, pI. 327), Lobosphinctes Buck cally successive, apomorphic structures, for simplicity man [M] (1923; type species: L. intersertus Buckman, 1923 respectively characterized by: (1) straight ribs of variable in 1909-1930, TA-V, pI. 447) and Phaulozigzag Buckman strength; (2) sigmoid ribs of variable strength and spac [m] (1926 in 1909-1930; type species: Phaulozigzag phaul- ing; and (3) distant, sharp, parabolic ribs. FIG. 3. Proc TAB LE 1. Gmer. of Zij;zagiceratmae mmtion Pro"rit<, Siemindzki. 1898 [type. Proc omorphus Buckman, 1926 in 1909-1930, TA-VI, pI. 643) the Meulam Formation (Lower Bathonian, Zigzag Zone) lad:rigzag-stadium and display complex suture lines. Pro also from G",at Balkhan ridges, and six sp"cimens of ",rites Siemiradzki [M] (1898, p. 78; type sp"cies : p, procc Pr"nchi" aff. qucll5tedti from 1he Upper part of M.ulam ru5 Schloenbach, 1865, by subsequent designation of Formation (also Zig>;ag Zone; in Bexnosov and Mitta Buckman 1914 and Opinion 301 of the International 1993, pI. 13, fig. 6). Beznosov wrote (in Bexnosov and Commission on Zoological Nomendatu",) and Sicmintdz Mitta 1993, p. 113) that Sibirjalova had not selected a ki" Hyatt [m] (1900, p. 582; typt species: Ammonites bakc holotype and he designated as lectotypt the specimen ri"c d'Orbigny, 1847 in 1842-1851, p. 424, pI. 149, fig. I) from Sibirjalova, 1961, pI. 6, fig. 2. Also, Beznosov show:rigzag-state I and a more complex suture line. noted that the indicated picture of Quenstedt (1886, pI. Sibirjalova (1961, p. SO, pl 6, figs 1-2) described a 64, fig. 9) is a rep",sentative of the Lower Bajocian new species Pmccrites qucll5tedti on the basis of two genus Emi!ci". Later, Mitta (in Beznosov and Mitta specimens from the 'Upp"r Bathonian sandstones' of 2000, p. 61) ",marked that, according to the Interna G",at Balkhan (TUlunenistan), but ",fened this species tional Code (International Commission on Zoological to Ammonites gc",inci gntndis Quenstedt (1886 in 1886- Nomendatu", 1999), 1he indication of Sibirjalova 1887, p. 514, pI. 64, fig. 9 only, not figs 4-8, 10-12). (1961) is valid as holotype identification for Quenstedt's Later, Beznosov described Pmnchi" qucll5tedti (Sibirjal specimen (i.e. Emi/ei" qucll5tcdti (Sibirjalova)), so that ova; in Beznosov and Mitta 1993, p. 112) on the basis the species p, qucn,tedti is subsumed in the n"" name of five specimens (I specimen figured in Beznosov and Pr"nchi" ,ibirj"""""c, with holotypt designated by Sibir Mitta 1993, pI. 13, fig. I), from the Lo"",r Member of jalova (1%1, pI. 6, fig. 2). Therefore, the 'Upp"r Batho- niiUl' syntypes of p, sibirju(YYu are not jwtifi.d ... eul.y Occurr fiG. 4. Franoh;" ar. .1U Stun"i; .,rompltte maerC>COl:lch; halatyp<, 'p Type specimen. The holotype PUll1399 [M] figured by Pavia Diagnosis. Evolute Franchia (U/D generally surpass et al. (200S, pI. 3, fig. 13) is refigured here in Figure 6A, from 45 per cent in post juvenile stages), macro- and micro- level BAll in Sturani (1967). conchs, with coarse ribbing. Type horizon. Lower Bathonian, Zigzag Zone, Macrescens Description. Adult shells of small to medium size, from micro Subzone. Level BAll of the 'Marno-calcaires a Cancellophycus' conchs surpassing 40 mm of diameter (Fig. 5D) to adult macro Formation in Bas Auran area (Sturani 1967) and level RA031 of conchs reaching 260 mm (Fig. 4). No specimens are known Ravin d'Auran Section (Pavia et al. 200S), indicated in Figure 2. possessing complete body ch amber, but umbilical suture sur passes 360 degrees in the macro conch holotype and 210 degrees Type locality. Bas Auran, AIpes de Haute Provence, south-east in the incomplete, microconch topotype (Fig. 7A). Evolute coil France. ing, with values of umbilical ratio ranging from 49 to 42 per cent, decreases in successive stages of ontogenic development (Fig. S), except by egression of umbilical seam in adult body Material. Three syntypes from the Macrescens Subzone of Bas chamber (Fig. 4B). Whorls vary in section from low-oval to sub Auran area. The paratype PU111573 [m] figured by Pavia et al. circular and high-oval contour, with convex flanks (Fig. 9A-D). (200S, pI. 3, figs 11, 11) and Fernandez-Lopez et al. 2009a, fig. Zigzag-state 2 developed in phragmocone, with sigmoid, sharp 6b) is refigured here in Figure 6B-C, magnified in Figure 7B-C, ribs of variable spacing (Figs 5C, 7A). Ulterior ornamentation to from level 11 in Sturani (1967) and RA033 in Pavia et al. zigzag-state consists of relatively coarse, straight to slightly sinu (200S), Ravin d'Auran Section. The paratype PU11157S [M], a ous, rounded ribs. Primary ribs usually bifurcate, with additional fragmentary nucleus c. SO mm in diameter, shows zigzag free intercalatories that pass over the venter radially or mth a stadium in the right side, up to 30 mm D and 20 mm U. gentle forward inclination. There are about 10-22 primaries per half whorl. Generally, secondary ribs are not interrupted on the Measurements. See Fernandez-Lopez and Pavia (2013). c Diagnosis. Moderately involute Franchia (U/D generally Type level. Lower Bathonian, Zigzag Zone, Macrescens Subzone. <45 per cent in post juvenile stages), macro- and micro Level BA9 of the 'Marno-calcaires a Cancellophycus' Formation conchs, with rounded and blunt ribbing. in Bas Auran area (Sturani 1967) and level RA024 of Ravin d'Auran Section (Pavia et al. 2008), indicated in Figure 2. Description. Adult shells of small to medium size, from micro conchs surpassing 35 mm of diameter (Fig. 6B-C) to macro Type locality. Bas Auran. conchs reaching 140 mm of fully septate phragmocone and expected to surpass 250 mm in diameter (Fig. 8). Only one Type region. AIpes de Haute Provence (Bas Auran and Chau lappetted microconch known (Fig. 7B-C), with complete body don), south-east France. chamber and umbilical egression but extremely short body chamber (130 degrees) indicative of preadult development. Diagnosis. Micro- and macrownchs of small or medium Moderately evolute coiling, with values of umbilical ratio rang size (40--300 mm). Planorbicones or discocones of mod ing from 41 to 48 per cent, decreasing in successive ontogenic erately evolute to involute coiling. Zigzag-state 2 devel stages (Fig. 8B). Whorls vary in section from low-oval to subcir oped in the inner whorls, but surpassing 10 mm in cular and high-oval contour, with convex flanks (Fig. 9E-F). diameter, firstly composed of proradiate to sub radiate ribs Zigzag-state 2 developed in phragmocone, with sigmoid, sharp of variable strength and spacing; then appear proradiate ribs of variable spacing (Fig. 7C). Ulterior ornamentation to zig zag-state consists of relatively coarse, straight to slightly sinuous, to sub radiate, sigmoid ribs of greater strength upflank rounded and blunt ribs (Fig. 6A). Primary ribs bifurcate or tri and variable spacing; and, finally, occasional proradiate to furcate, with additional free intercalatories that pass over the sub radiate, sharp parabolic ribs of variable spacing. venter radially or with a gentle fo nvard inclination, without Numerous, blunt, bifurcate or trifurcate costae. Relatively interruption. There are about 10-19 primaries per half whorl wmplex suture line, with moderately broad andlor rela Looped, weak ribs, united on ventrolateral angle at a parabolic tively short lateral lobe. node, occur in the outer whorl of the microconch (Fig. 7C). At maturity, ribs become sparser and blunter on flanks, and eventu Remarks. Sturani (1967) regarded Z. torrensi and Z. tor ally fade out, while still persisting on the venter. Shallow, broad rensi variecostatum as microconchs of dimorphic macro and prorsiradiate constrictions are present in the holotype. wnchs from the upper Macrescens Subzone, such as Suture line relatively simple, with suspensive lobe not strongly Zigzagiceras (Procerozigzag) postpoIIubrum Wetzel garnieri. retracted. However, Zigzagiceras Buckman [m] (1902, p. 7; type spe cies: Ammonites zigzag d'Orbigny, 1846 in 1842-1851, p. Remarks. Franchia arkeIli Sturani, the type species of the 390, pI. 129, figs 9, 10) and Procerozigzag Arkell [M] genus, is the most similar representative. F. subalpina, (1953, p. 37; type species: Stephanoceras crassizigzag Buck however, shows stouter whorls and more involute coiling. man, 1892, pI. 14, figs 2, 3) display zigzag-state 3 and These two species are chronostratigraphically coincident more complex suture lines. The distant parabolic ribs of at the basal Macrescens Subzone, but F. subalpina persist Zigzagiceras [m & M] become strongly rursiradiate on into younger intervals, whereas F. arkelli occurs at the lower and middle flanks, sharply projected forward on top of the precedent Parvum Subzone. the ventrolateral shoulder, with parabolic tubercles or nodes, and the septal suture is strongly retracted with Occurrence. The syntypes of F. subalpina correspond to the Macrescens Subzone of the Bas Auran area. It is an endemic spe very narrow and long lateral lobe. Comparatively, Proto & cies of the Submediterranean Province, in Subalpine areas near zigzagiceras [m M] is a lasting phyletic lineage of con the Provence-Ardeche platform system, in the northern border siderable morphological variability, and the source of the of the western Tethys, phyletically derived from F. arkelli. more specialized and briefer Zigzagiceras [m & M]. Both macroconchs and microconchs of Franchia also display zigzag-state 2, but show a simpler suture line, more Genus PROTOZIGZAGICERAS Fernandez-Lopez et al. 2007 planorbiconic coiling, shorter chronostratigraphical range and more restricted palaeogeographical distribution. Type species. Zigzagiceras torrensi (Sturani 1967, p. 47, pl 21, Zigzagiceras Ienthayense [m] sensu Sturani (1967, p. 48, fig. 3), by original designation of Fernandez-Lopez et al. 2007, pI. 20, fig. 4) non Arkell (1958 in 1951-1958, p. 204, pI. p. 396. 23, fig. 6a-b), from the level BA8, upper Macrescens FIG. 5. Franchia arkelli Sturani. A-C, incomplete phragmocone of macroconch; paratype, specimen PU3 1691, level BAl in Sturani (1967), Le Touert Section, Chaudon. D, complete microconch, topotype, specimen PU112318, level RR033 in Pavia et al. (2008) and level BAll in Sturani (1967), Ravin des Robins Section, Bas Auran; for magnified view, see Figure 7A. E-F, incomplete phragmocone of macroconch; paratype, specimen PU31690, level BAl in Sturani (1967), Le Touert Section, Chaudon. All specimens from the Macrescens Subzone, Zigzag Zone. Black asterisk marks the last septum of the phragmocone. Scale bars represent 10 mm. , :\, .�, � , '' ." . , ,.t , I I",i, 't' , ;,.l v ' .\ " , , FIG. o. Franchia 'Wo A. Franch", ,ubdpina 'P' nov.; incomplete phr.gmocone of m.c:roconch; halotyp<. 'pecIneJl PUll1399. level RA030 ., Pavi. 'Ial (2008) and level BAll in Sturmi (1967). B-C, F ,ubalp;"" 'po nov.; complete microconch; paratJP<. 'p SUbzone, and Pmcrrite5 imitator Buclman [M] in Sturani 12, fig. 1) and lrm (�afu.deh md Seyed·Emami 2005, fip 10, {1967, p. 48, pI. 15, fig. 5} and Torrens {1987, pI. 7}, 11; Sh.Jru md Sey";·Em.mi 2010, p. 2, fig. 12; T.ble 2). Judg· from the levels BAI to BA6, Aurigerus Zone, also display mg by the>e referme<>, ProrozilP"'tx"a, Occurs in the Early :rigzag-state 2, but art mort planorbiconic and of larger B.thonim, from Parvum to Recinctrn subzone>, Mediterrane.n adult size. On the other hand, Zigz "gicu -" 20 ·-4.- Zigzagiceras (Procerozigzag) postpollubrum Wetzel garnieri figured . .... , . ._.- '- by Sturani (1967, pI. 21, fig. 7a-b) and refigured here in ...... - . .,;..._ -- 10 Figure 11, and PU3ll72S [M] collected by Sturani in 1972 and o figured by Torrens (1987, pI. 9, fig. 3a-b). One specimen from 40 50 60 70 80 90 100 110 120 130 140 150 160 mm level BA8 or BA9 collected by Sturani in 1972, PU31720 [M], B and figured in Figure 12. One specimen from level BA9 or BA10 80 mm collected by Sturani (1967), PU311711 [M], and figured by Tor u 60% rens (1987, pI. 8, fig. 2a-b). One specimen from level BAll in 70 • Sturani (1967) and RA031 in Pavia et al. (2008), PUll1S77 [M], 60 figured by Pavia et al. (2008, pI. 3, figs 9, 10). 50 ... 30% Syntypes. There are two syntypes. The paratype PU31677 [M] 40 • •• described but not figured by Sturani (i 967, p. 48) is figured here 30 in Figure IOI-J. 20 New material. Four topotypes: PU31680 [m] figured III 10 III 0 Figure IOG-H; PU31681 [m] in Figure IOE-F; PU31682 [m] Figure 10K-M; and PU31720 [m] in Figure 12. 40 50 60 70 80 90 100 110 120 130 140 150 160 mm C Measurements. See Fernandez-Lopez and Pavia (2013). 140 W/H • 130 Diagnosis. Protozigzagiceras, planorbicones to discocones 120 • • of moderately evolute to involute coiling, with straight to • .... . slightly concave, rounded and blunt, generally bifurcate ribs. 110 • *h 100 • • • 11> h. Description. Adult shells of small to medium size, from incom • .& 90 h • plete micro conchs reaching 70 mm diameter (Fig. lOG-H) to 0 macroconchs reaching 2S0 mm in diameter. Moderately evolute 40 50 60 70 80 90 100 110 120 130 140 150 160 mm coiling, with values of umbilical ratio ranging from 49 to 41 per cent, decreasing insuccessive stages of ontogenic development D (Fig. 13B). Whorls vary in section from low-oval to subcircular 25 Ni/2 and high-oval, with convex flanks (Fig. 14A-D). Zigzag-state 2 •• 20 • developed in the inner whorls, composed of proradiate to subra 15 I it.. t;r • diate, sigmoid ribs of greater strength upflank and variable spac ing; and, finally, occasional, sharp parabolic ribs of variable 10 - .:rt, . *. �h h spacing (Fig. IOD). Ulterior ornamentation consists of straight u to slightly concave, rounded and blunt, generally bifurcate ribs 10 20 30 40 50 60 70 80 90 100 110 120 130 mm that cross the venter with a gentle forward inclination. Simple primaries and intercalatories occur. There are about 19-26 FIG. 8. Plots of measurements of Franchia and Megazigzagi primaries per half whorl. Generally, secondary ribs not inter A, whorl width (W) versus shell diameter (D). B, umbilical ceras. rupted on the middle of the venter. Shallow and prorsiradiate diameter (U) versus shell diameter (D). C, whorl width/whorl constrictions present exceptionally (Fig. lOB, J). Relatively com height ratio (W/H) versus shell diameter (D). D, internal ribs per plex suture line, with relatively short lateral lobe (Fig. 15). half whorl (Nil2) versus umbilical diameter (U). h, holotype. Remarks. According to Sturani (1967), z. torrensi is the Type horizon. Lower Bathonian, Zigzag Zone, Macrescens Sub zone. Level BA9 of the 'Marno-calcaires a Cancellophycus' Forma micro conch counterpart of Procerozigzag postpollubrum tion in Bas Auran area (Sturani, 1967) and level RA024 of Ravin garmen, and Zigzagiceras postpollubrum appears to be d'Auran Section (Pavia et al. 2008), indicated in Figure 2. the end member of a wide chronospecies in which the . . # ...... • # . . • # . . . . • # . . f. ' .. lr #. le . . . :: . . . . # • ...... • • • 00c D E B FIG. 9. Whorl shape cross-sections, through the phragmocone and body chamber (stippled), of Franchia spp. A-D, Franchia arkelli Sturani; A, holotype, PU31689 [M] (Fig. 4); B, paratype, PU31690 [M] (Fig. SE-F); C, topotype, PU11132S [M] (Fig. 6D); D, para type, PU31691 [M] (Fig. SA-C). E-F, Franchia subalpina sp. nov.; E, holotype, PUl11399 [M] (Fig. 6A); F, paratype, PUl l1S73 [111] (Figs 6B-C, 7B-C). Scale bar represents 10 111111. maximum size of full-grown specimens reaches 250 mm. pI. 19, fig. 5) was based on four specimens from the beds In Bas Auran (Sturani 1967, pI. 22, fig. 1; Torrens 1987, succeeding that yielding the holotype of P. torrensi (BA8 pI. 8, figs 3-5), Protozigzagiceras postpollubrum (Wetzel and BA7). They show the zigzag-stage more densely 1937, p. 107, pI. 11, fig. 3a-b) comprises involute disco ribbed and proportionally shorter, ending at 10 mm. On cones with a very narrow and deep umbilicus «24 per account of their biochronostratigraphical and morphological cent in adults), and with an earlier fade of primaries and differences, this subspecies was regarded as transitional secondaries at 80 and 120 mm, respectively, at the top of between typical Zigzagiceras and typical Siemiradzkia. the Macrescens Subzone (level BA7 in Sturani and level However, the holotype is lost (H. Torrens, pers. comm.), RA019 in Pavia et al. 2008). the paratypes are fragmentary inner whorls, and new Zigzagiceras torrensi variecostatum Sturani (1967, p. 48, specimens similar to the holotype are unknown at pI. 20, fig. 2a-b [holotype], pI. 2, fig. 5; pI. 13, fig. 4a,c; present. TABLE 2. Specimens published belonging to the genus Proto fig. 10) is refigured here in Figure 16H-I; both specimens from zigzagiceras Fernandez-Lopez et al. 2007. level BA13 in Sturani (1967) and RA035 in Pavia et al. (2008), Protozigzagiceras Biohorizon, Parvum Subzone. The paratypes References Occurrence PU31695 [m] figured in Figure 16A-D and PUll1299 [m] fig Buckman (1892, pI. 13, figs 1, 2) England ured in Figure 16E-G from level BA12 in Sturani (1967) and RA033 in Pavia (2008); Macrescens Subzone. Two frag Wetzel (1937, pI. 11, fig. 3) France et al. Sturani (1967, pI. 13, fig. 4, pI. 15, fig. 1, pI. 19, France mentary paratypes, PUllI572 [m] and PUll 1300 [m], respec fig. 5, pI. 21, figs 1, 3, 7, pI. 22, fig. 1) tively, from levels BA12 (= RA033) and RA035. Galacz (1980, pI. 32) Hungary Sandoval (1983, pI. 35, fig. 3) Spain Measurements. See Fernandez-Lopez and Pavia (2013). Torrens (1987, pI. 8, figs 2-5) France Dietze and Chandler (1997, pI. 1, figs 3, 4) England Diagnosis. Protozigzagiceras, slim planorbicones of moder Schlogl et al. (2005, pI. 10, figs 4, 5, pI. 12, fig. 1) Slovakia ately evolute to moderately involute coiling, with straight, Shafizadeh and Seyed-Emami (2005, figs 10, 11) Iran rounded and blunt, generally bifurcate ribs. Fernandez-Lopez et al. (2007, fig. 11) Portugal Fernandez-Lopez et al. (2007, figs 10, 12) France Description. Fully septate, discoid shells of small to medium size, Pavia et al. (2008, pI. 3, figs 6, 7, 9-12) France from microconchs reaching 50 mm diameter (Fig. 16E-G) to pre Fernandez-Lopez et al. (2009a, fig. 6b) France sumed macroconchs surpassing 60 mm (Fig. 16H-I). Moderately Shams and Seyed-Emami (2010, p. 2, fig. 12) Iran evolute coiling, with values of umbilical ratio ranging from 49 to 39 per cent, decreases in successive stages of ontogenic develop Occurrence. Protozigzagiceras torrensi is known from the French ment (Fig. 13B). Whorls vary in section from low-oval to subcir Subalpine Basin, restricted to the Macrescens Subzone. It is com cular and high-oval, with convex flanks (Fig. 14E-H). Zigzag-state mon in the upper part of this subzone and scarce in the lower part. 2 developed in the inner whorls, firstly composed of proradiate to All the microconchs of Figure 10 come from the level BA9 in subradiate ribs of variable strength and spacing; then appear pro Sturani (1967) and RA029 in Pavia et al. (2008), Ravin d'Auran radiate to subradiate, sigmoid ribs of greater strength upflank and Section, but the macroconch published by Pavia et al. (2008, pI. 3, variable spacing (Fig. 16G). Ulterior ornamentation consists of figs 9, 10) corresponds to the level RA031 (= Sturani's level BAll). straight to slightly concave, rounded and blunt, generally bifurcate ribs that cross the venter radially or with a gentle forward inclina tion. Simple primaries and intercalatories occur. There are about 19-23 primaries per half whorl. Generally, secondaries are not Protozigzagiceras tethycum sp. novo interrupted on the middle of the venter. Shallow and prorsiradiate Figures 13, 14E-H, 16 pseudoconstrictions or constrictions are very scarce. LSID. urn:lsid:zoobankorg:act:A077DDEE-E86A-4AE4-AA76- Remarks. Among the syntypes, the microconchs show a 6666D62B9A73 zigzag-state 2 clearly longer than the macroconch (Fig. 16H-I) as in other zigzagiceratin species. Protozigzagiceras 2007 Protozigzagiceras g. novo sp. aff. P. torrensi (Stura ni); Fernandez-Lopez et aI., p. 396, fig. 10. tethycum comprises forms more planorbiconic and appar ently smaller in size than P. torrensi. It differs from Fran chia [M & m] by the more complex suture line, whereas Derivation of name. After the Neo-Tethys Ocean and the Peri Siemiradzkia [m] lacks zigzag-state 2 with sigmoid ribs. Tethyan sedimentary basins, where the species has been com Planisphinctes [m] - Lobosphinctes [M] and Phaulozigzag monly recorded. [M & m] lack zigzag-stadium. Type specimen. The holotype PUlll574 [m] is figured in Figure 16J-L. Occurrence. The holotype of P. tethycum corresponds to the Proto zigzagiceras Biohorizon, upper Parvum Subzone,but other syntypes come from the lowermost Macrescens Subzone, Bas Auran area. Type horizon. Lower Bathonian, Zigzag Zone, Parvum Subzone, Protozigzagiceras Biohorizon. Level BA13 of the 'Marno-calcaires a Cancellophycus' Formation in Bas Auran area (Sturani 1967) and level RA035 ofRavin d'Auran (Pavia d al. 2(08), indicated in Figure 2. Protozigzagiceras flexum sp. novo Figures 13, 17 Type locality. Bas Auran, AIpes de Haute Provence, south-east France. LSID. urn:lsid:zoobankorg:act:9E99CI21-3A32-40E3-AB2F- 665B256C76C2 MateriaL Six syntypes, all fully septate phragmocones, from the Parvum and Macrescens subzones of Bas Auran area. The para Derivation of name. After the flexuous ribs that characterize the type PU31694 [M] figured by Fernandez-Lopez et al. (2007, species. - A , FIG. , O. hotozigzap"ra, /onon,; (Sturmi). A-D. incomplete phragmocon<; holotyp<. sp Ty p. 'p",mm The holot)'J'< PUll1326 [m] i> figured in Ty p. horizon. lower BathQnian. Zi17ag Zone, Macre>c FIG. , ,. i'roloziIP'P"ra, /onon,; (Sturmi); inrompl Ty p. locdity BM Amm, AIr de Haute Proven"" <""th-eMt France. Matorid Two M"" ur Dia.gnosi!;. Pmtazigza.giccrct5, planorbicones to discocones of moderately "volute to moderately involute coiling, with flexuous, rounded and blunt, generilly bifurcate ribs. FIG. '2. horozigzap"ra, torrond (Sturani); incompl D - 37 per cmt), wih COn"", flanh. Zigzag- 10 the venter. D Remarks. Protozigzagiceras flexum is close to P. tethycum, 40 50 60 70 80 90 100 1110 120 130 140 1150 160 mm in its moderately evolute coiling, but differs in the more B 80 mm depressed whorl section and the flexuous ribs with U . 55% forward inclination on the ventral region, instead straight 70 .... .- ribs crossing the venter radially. 60 '" .,. Zigzagites Buckman [M & m] (1922; type species: Zig .- 50 zagites imitator Buckman, 1922 in 1909-1930, TA-IV, pI. � 31}% 301), Zigzagiceras lenthayense [m] in Sturani (1967, p. 48, 40 pI. 20, fig. 4) non Arkell (1958 in 195 1-1958, p. 204, pI. 30 23, fig. 6a-b), from the upper Macrescens Subzone, Zig 20 zag Zone, and Procerites imitator Buckman [M] in Sturani 10 (1967, p. 48, pI. 15, fig. 5) and Torrens (1987, pI. 7) from e" D the Aurigerus Zone, also display zigzag-state 2, but the 40 50 60 70 80 90 100 1,10 120 130 140 1150 1160 mm phragmocone is more plan orbiconic and of larger adult C size. However, they could be counterpart dimorphs and 140 W/H paedomorphic taxa originated from a more involute and h 130 "'6 depressed species such as P. flexum during the Macrescens Subzone. Analogously, Wagnericeras could be a paedo 120 .t:. 6 morphic genus originating from this group of flexicostate 110 e" .... zigzagiceratins during the Macrescens Subzone, taking .... � e" 100 .6 e" into account of the relatively evolute specimen identified 6 90 0 by Torrens (1987, p. 46, pI. 20, fig. 1) as Wagnericeras? eh D sp. nov., collected by Sturani in bed BA8, and probably 40 50 60 70 80 90 100 110 120, 130 140 150 160 1'111'11 the earliest known representative of that genus. D Occurrence. Protozigzagiceras flexum is very scarce in the lower 25 Ni� most Macrescens Zone of Bas Auran, probably derived of P. tethy 20 Me" cum at the latest Parvum Subzone (Protozigzagiceras Biohorizon). he" 15 .... h 10 .... U Protozigzagiceras densum sp. novo Figures 13, 18 10 20 30 40 50 60 70, 80 90 100 110 120 130 rmm FIG. 1 3. Plots of measurements of Protozigzagiceras. A, whorl L51D. urn:lsid:zoobank.org:act:5656C765-CEE3-45Al-B82E- width (W) versus shell diameter (D). B, umbilical diameter (U) 6FABA2AB4B8C versus shell diameter (D). C, whorl width/whorl height ratio (W/H) versus shell diameter (D). D, internal ribs per half whorl 2008 Franchia [m 1 arkelli Sturani; Pavia et aI., pI. 3, (Ni/2) versus umbilical diameter (U). h, holotype. figs 6, 7. developed in the inner whorls, up to 25 mm in diameter, with proradiate to subradiate, sharp sigmoid ribs of greater strength Derivation of name. After the dense ribbing that characterizes upflank and variable spacing; and, later, numerous, slightly con- the species...... , , . , <. : > : . . 9 " . , I' " I' ' , . .. ,. " .. 'to .f -..,. ".t " .. , 'to" ... 0" ,. ;t @" '@ Q . , . , . , I be bD OF OG0 o. 8E FIG. 14. Whorl shape cross-sections, through the phragmocone and body chamber (stippled), of Protozigzagiceras spp. [m & M]. A-D, Protozigzagiceras torrensi (Sturani); A, paratype, PU31677 [M?] (Fig. IOT-J); B, topotype, PU31681 [M] (Fig. IOE-F); C, topo type, PU3l682 [m] (Fig. IOK-M); D, holotype, PU3l676 [M] (Fig. lOA-D). E-H, Protozigzagiceras tethycum sp. nov.; E, paratype, PU3l694 [M] (Fig. l6H-I); F, paratype, PUl11299 [m] (Fig. l6E-G); G, paratype, PU l11574 [m] (Fig. l6J-L); H, holotype, PU3l695 [m] (Fig. l6A-D). Scale bar represents 10 mm. Description. The holotype is a fully septate phragmocone of mi croconch (Fig. l8A-D) surpassing 40 mm in diameter and with moderately involute coiling, with values of umbilical ratio c. 45 per cent (Fig. l3B). Whorl section low-oval (W/H = 1.2 and W/D = 39 per cent), with convex flanks. Ornamentation, mainly ribbing and parabolic nodes, is very variable. Zigzag-state 2 developed in the inner whorls, but surpassing 30 mm in diam eter, with proradiate to subradiate, sharp sigmoid ribs of greater strength up flank and variable spacing; and, later, occasional pro radiate to subradiate parabolic ribs of variable spacing. Last half whorl displays numerous, straight to slightly concave, rounded and blunt, subradiate primary ribs (Nil2 = 20). Secondary ribs, FIG. 15. Septal suture of Protozigzagiceras torrensi (Sturani), bifurcate or trifurcate, are quite weak, wiry, crossing the venter specimen PU31682 (Fig. IOK-M). E, external lobe; L, lateral with forward inclination (Ne/2 = 53, i = 2.6). Whorl thickness lobe; U2, second umbilical lobe. Scale bar represents 10 mm. is at its maximum just at the ventrolateral margin, where com mon parabolic nodes occur in the last preserved whorl. Shallow and prorsiradiate constrictions are locally present. Relatively Type specimen. The holotype PUll1322 [m], figured by Pavia complex suture line. et al. (2008, pI. 3, figs 6, 7) and refigured here in Figure l8A-D. Remarks. Protozigzagiceras densum is the most involute, Type horizon. Lower Bathonian, Zigzag Zone, Macrescens Subz deeply navel and early polyfurcate species of the genus one. Level BA12 of the 'Marno-calcaires it Cancellophycus' For mation in Bas Auran area (Sturani 1967) and level RA033 of recorded in Bas Auran area at the transition between Par Ravin d'Auran Section (Pavia et al. 2008), indicated in Figure 2. vum and Macrescens subzones. Moreover, the holotype is one of the specimens of this genus showing the largest Type locality. Bas Auran, Alpes de Haute Provence, south-east number of parabolic nodes. This species clearly differs France. from species of Zigzagiceras [m] - Procerozigzag [M] in lacking zigzag-state 3, with distant and sharp, parabolic Material. One specimen, the holotype PU l11322 [m], Fig ribs. Another specimen, showing early polyfurcate ribs ure 18A-D. ventrally projected forwards, with zigzag-state 2 formed of sharp, sigmoid and parabolic ribs of variable spacing, Measurements. See Fernandez-Lopez and Pavia (2013). has been figured from the Protozigzagiceras Biohorizon of Cap Mondego by Fernandez-Lopez et al. (2007, fig. 11). Diagnosis. Protozigzagiceras, planorbicones to discocones That Portuguese specimen is more planorbiconic and of moderately involute coiling, with straight to slightly older than the holotype of P. densum, but it can represent sinuous, rounded and blunt, bifurcate and polyfurcate ribs. a plesiomorph of the same phyletic lineage. •• G FIG. , {;. hotozigzagi"ra, t,thycwn 'po nOvo A-D, inCOITlJx The two Cbilean 'p"cimens taxonomically determined Sierra de Argomedo, Domeyko Cordillera, Lower Batho ?Zigza.gicuct5 sp. (Gro.chh ""cl VDn HiIlebrandt 1994, pI. nian, correspond to immature or preadult, fragmentary I, figs 4, 5; Groschh 1996) from Aguada El Om and W shells. They display sigmoid ribs of variable strength and A FIG. , 1. Protozigzagi"ra, fioxum 'po nOVoA-D, incQmp. splcing, bifurcate or trifurcate that Cto.. the venter Ty p. toediry Dhmnl uel, •."tnl Saudi Anbi.•. ndiilly (therefore zigzag-stm 2). v.r;...�tWn ofn�",. Aft ..'he lugc .u. ...dled by Ihe odult. of the tl'P'"' ","" . 0""",,,,,, P'M:ip(ial�, Wuk'" i.< 0 -r .cuor: >peci'" in the lowtnnG>1 MaaCOG Occurron", M,tazigzag>; FIG. , 9. M,tazpgi"ra, ,ubarabicum, gm. d 1he relatively developed 2ip;ag-state in 1he phragmocone of Ty p. !ocauly, W.di al Hisy;m (24"45'N), Dharn,. Qu.drmgl, 1hese two evolute, planorbiconic holotypes, bo1h are mi (cmtral S.udi Ar.bia). croconchs. Bo1h species of Epizigzagicercrs are associated wi1h CobbMites mlkeetnanUl" lmlay, referred to Late Bajo Mat Cl) Cl) Early Bathonian � Cl) � Cl) Cl) � Zigzagiceratinae Cl) .� Cl) Cl .� Co) � co co Cl) .8J Co) .� .� co .8J ..c:: .�N N N co Co) co .� c: .8 Cl .� Biochoremas � � e Cl) N IJ.. Q � tfr Tethys-Panthalassa Realm : Mediterranean-Caucasian Subrealm ... * • Indo-Pacific Subrealm * East-Pacific Subrealm 7'" + Zigzagiceras ... + Epizigzagiceras Franchia * Arcto-Pacific Subrealm + * Megazigz agiceras Protozigzagiceras • FIG. 20. Palaeobiogeographical distribution of some Zigzagiceratinae genera during Early Bathonian Zigzag Zone. Palaeogeography modified from Fernandez-Lopez et al. (2009b), Fernandez-Lopez and Chong Diaz (2011) and Golonka (2011); palaeobiogeographical units according to Westermann (2000b). other hand, Megazigzagiceras [m] belongs to the Indo Late Bajocian Early Bathonian Pacific Subrealm (Ethiopian Province; Enay 2007), Parkinsoni Zigzag Aurigerus whereas Epizigzagiceras [m & M?] characterizes the East G) '" 0 ;;: col CD OJ ""0 III ;0 ::J Pacific and Arcto-Pacific subrealms (Athabascan and Ber DJ ::J 0 III " CD c: ;:a. » Stephanov (1972), this dimorphic group can be included FIG. 21. Phylogenetic scheme of the Early Bathonian genera of among the youngest Leptosphinctinae (Westermann 1956, Zigzagiceratinae, originated from Leptosphinctinae and diversi 1958; Luppov and Druzschic 1958; Beznosov and Mikhail tied during Parvum and Macrescens subzones (Zigzag Zone, ova 1981; Beznosov 1982; Torrens 1987; Innocenti et al. Early Bathonian), before the extinction of Bigotites. Geochrono 1990; Beznosov and Mitta 1993, 1995, 1998; Fernandez logical divisions after Pavia et al. (2008) and Fernandez-Lopez Lopez et al. 2006, 2007). The phylogenetic derivation of et al. (2009a, b). Zigzagiceratinae, in particular Zigzagiceras [m] - Procero zigzag [M] and Franchia [M & m] , from Bigotites [M & m] whereas Zigzagiceras [m] - Procerozigzag [M] represents a as supported by Sturani (1967) and Torrens (1987), or neotenic zigzagiceratin branched off from Protozigzagi from Phaulozigzag [m] - Lobosphinctes? [M] as suggested ceras [m & M]. P. tethycum and F. arkelli appear progenetic by Fernandez-Lopez et al. (2006), remains to be reassessed because they attain maturity at a relatively small size. according to very accurate biochronological data and com Protozigzagiceras [m & M] and Franchia [M & m], besides plementary morphological information (Fernandez-Lopez zigzag-state 2, display morphological features characteris et al. 2006, 2007; Pavia et al. 2008). tic of the earliest Procerites at a juvenile stage, such as rel Procerites [M] - Siemiradzkia [m], the oldest known atively planorbiconic and evolute coiling with relatively Zigzagiceratinae with zigzag-state 1, probably branched simple suture lines. Protozigzagiceras [m & M] spreads in off from the alleged Leptosphinctinae Phaulozigzag [m] - middle latitudes to many northern Peri-Tethyan basins, Lobosphinctes? [M] group (Fig. 21). The holotype of from Proto-Atlantic Ocean and Lusitanian Basin (Buck P. phaulomorphus Buckman (1926 in 1909-1930, TA-VI, man 1892, pI. 13, figs 1, 2; Dietze and Chandler 1997, pI. pI. 643) lacks a zigzag-stadium, but is a very small adult 1, figs 3, 4; Fernandez-Lopez et al. 2007, fig. 11), through (D = 56 mm), and the possible occurrence of parabolae Alpine Ocean, Subalpine Basin and Tizia (Sturani 1967, in this taxonomic group needs further investigation. pI. 13, fig. 4, pI. 15, fig. 1, pI. 19, fig. 5, pI. 21, figs 1, 3, 7, Protozigzagiceras [m & M] and Franchia [M & m] rep pI. 22, fig. 1; Galacz 1980, pI. 32; Sandoval 1983, pI. 35, resent progenetic zigzagiceratins, directly or indirectly fig. 3; Torrens 1987, pI. 8, figs2-5; Schlogl et al. 2005, pI. branched off from Procerites [M] - Siemiradzkia [m] , 10, figs 4, 5, pI. 12, fig. 1; Fernandez-Lopez et al. 2007, figs 10, 12; Pavia et al. 2008, pI. 3, figs 6, 7, 9-12; Fernan Subwne (Fernandez-Lopez 2000, fig. 2, pI. 1, fig. 9). In dez-Lopez et al. 2009a, fig. 6b), to the Iranian platform turn, plesiomorphic representatives, older and more plan system (Shafizadeh and Seyed-Emami 2005, figs 10, 11; orbiconic than P. densum, have been recorded in the Shams and Seyed-Emami 2010). On the other hand, uppermost Parvum Subzone of Cap Mondego. Therefore, Franchia seems to be endemic to Subalpine areas, near most of the earliest Bathonian phyletic lineages of the the Provence-Ardeche platform system. clade Zigzagiceratinae developed lasting peramorphoclines Zigzagiceras [m] - Procerozigzag [M], in addition to with increasing hydrodynamic coiling of the shell and zigzag-state 3 ribbing and a complex suture line, displays increasing adult size (at least, in the microconchs), in an a relatively planorbiconic and evolute coiling, in particu opposite sense to the previous, rapid, paedomorphic lar Zigzagiceras euryodos Schmidt [m] (1846, p. 106, pI. changes. 43, fig. 6a-b) and Zigzagiceras pseudoprocerum Buckman In conclusion, the ancestor of the clade Zigzagiceratinae [M] (1926 in 1909-1930, TA-VI, pI. 623). This dimorphic can be found in the genus Procerites, rapidly diversified in group becomes widely distributed along the northern bor multiple phyletic lineages by iterative, paedomorphic der of Western Tethys. The primitive and evolute Zigzagi changes and additional, lasting, peramorphic modifications ceras, in particular Z. euryodos [m & M], may be a direct during the Early Bathonian Zigzag Zone. The Mediterra neotenic derivative of P. densum, at the basal Macrescens nean-Caucasian genera Franchia, Zigzagiceras, Zigzagites Subwne. The group of Z. imitator Buckman [M] (1922 and Wag nericeras are phyletically derived of successive spe in 1909-1930, TA-IV, pI. 301) and Z. lenthayense [m] cies of Protozigzagiceras, which is itself a direct derivative of sensu Sturani (1967, p. 48, pI. 20, fig. 4) non Arkell (1958 Procerites. This evolutionary pattern produced numerous in 1951-1958, p. 204, pI. 23, fig. 6a-b) was suggested as and brief phyletic lineages with common homeomor homeomorphic with Procerites by Torrens (1987). This phisms. These new palaeontological results concerning the group, and the stouter group of Wag nericeras Buckman oldest Zigzagiceratinae are of biochronostratigraphical [M & m] (1921 in 1909-1930, TA-Ill, p. 33), both importance for calibration of the basal Bathonian and cor recorded at the level BA8 (in Sturani 1967), may have relation of the Bathonian GSSP. originated by neoteny from more involute and depressed species such as P. flexum during the Macrescens Subzone. Acknowledgements. Thanks are due to the Reserve Naturelle In this way, Zigzagites [M & m] and Wag nericeras [M & Geologique de Haute-Provence (Digne-Ies-Bains, France) for m] could be two neotenic zigzagiceratins, independently permission to work and sampling of the Bas Auran sections. We express thanks to R. Enay (Claude-Bernard University, Lyon, branched off from Protozigzagiceras. Also, Megazigzagi France) for making the specimen from Dharma, Saudi Arabia, ceras and Epizigzagiceras, showing zigzag-state 2 and lack of Figure 19, available for study, and to V. V. Mitta (Borissiak ing distant, sharp, parabolic ribs, can be regarded as Paleontological Institute, Moscow, Russia) for complementary neotenic groups phyletically derived from diverse species information on the alleged Caucasian species of Franchia. The of Protozigzagiceras. Both derivative genera probably cor authors also thank Kevin Page (School of Earth, Ocean and respond to Macrescens and/or post-Macrescens intervals, Environmental Sciences, University of Plymouth, England), an being very improbable from any Late Bajocian interval anonymous referee, Svend Stouge, Andrew Smith, Julia Sigwart (Fig. 21). The phyletic position of these two large-sized and Sally Thomas for constructive suggestions of the manu genera and the above-mentioned Chilean zigzagiceratins, script. This work was fu nded by the CGL2008-01273 (MICINN) however, still requires further investigation. and CGLlOll-23947 (MICINN) research projects (SRF-L) and In summary, rapid proterogenesis gave rise to the earli the Torino University Grants 2008 (GP). est Bathonian zigzagiceratin lineages, producing paedo morphic members, commonly neotenic and more scarcely progenetic (Fig. 21). In contrast, the successive species of DATA ARCHIVING STATEMENT Protozigzagiceras (from P. tethycum to P. torrensi and Data for this study are available in the Dryad Digital Repository: P. postpollubrum) and Franchia (from F. arkelli to F. sub http,!!dx.doi.orf!!l 0.s061/ dryad. gfli80 alpina) show trends towards increasing involution of ini tially evolute planorbicones to more weakly ornamented Editor. J ulia Sigwart discocones in the Bas Auran area. 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Fernandez-Lopez * and Giulio Pavia l Departamento de Paleontologia, Facultad de Ciencias Geol6gicas, calle Jose Antonio Novais 2, Universidad Complutense de Madrid, 28040-Madrid, Spain; e-mail: [email protected] 2 Dipartimento di Scienze della Terra, via Valperga Caluso 3S, IOI2S-Torino, Italy; e-mail: giulio.pavia@unito .it * Corresponding author Measurements A. Measurements for the sEecimens of Franchia arkelli Sturani. 1967 Specimens Figures D H HID W WID U UID WIH Ni/2 Nel2 Figs. 4A-B PU31689 [M] 220.0 60.0 27.3% 61.2 27.8%) 101.6 46.2% 1.02 14 35 2.5 hoiotype 170.0 53.8 31.6% 52.0 30.6% 78.2 46.0% 0.97 15 PU31690 Figs. 5E-F [M] 147.0 43.5 29.6% 40.0 27.2% 71.8 48.80/0 0.92 114.0 34.5 30.3% 34.7 30.4% 54.8 48.1% 1.01 20 45 2.3 PUl12317 [M] 92.0 31.9 34.7% 29.4 32.0% 40.2 43.7% 0.92 16 PU31691 Figs. SA-C [M] 70.0 22.5 32.1% 22.0 31.4% 33.0 47.1% 0.98 53.0 13.7 25.8%) 16.8 31.7% 23.9 45.1% 1.23 22 PUl11325 Fig. 6D [M] 75.0 23.8 31.7% 23.3 31.1% 31.9 42.5% 0.98 17 PUl12318 Figs. 4D, 7A [m] 31.5 9.3 29.5% 9.8 31.1% 14.6 46.3% 1.05 B. Measurements for the sEecirnens of Franchia subaZe.ina sE. novo Specimens Figures D H HID W WID U UID WIH Nil2 Nel2 Fig. 6A PUl11399 [M] 140.0 46.5 33.2% 43.0 30.7% 57.7 41.2% 0.92 45 hoiotype 75.0 26.0 34.7% 30.0 40.0% 33.0 44.0% 1.15 16 PUl11578 [M] 45.0 12.5 27.8%) 20.1 44.7% 19.2 42.7% 1.61 16 Figs.6B-C, 7B- PUl11573 [m] 32.0 10.5 32.8%) 9.8 30.6% 15.3 47.8%) 0.93 19 32 1.7 C C. Measurements for the specimens of Protozigzagiceras torrensi (Sturani) Specimens Figures D H HID W WID U UID Will Nil2 Nel2 Torrens 1987, PU31725 [M] 115.6 41.0 35.5% 45.1 39.0% 43.8 37.9%) 1.10 21 44 2.1 pI. 9, fig. 3 94.4 33.7 35.7% 38.0 40.3% 37.7 39.9%) 1.13 20 44 2.2 PU31717 Figs. 11A-B [M] 105.7 46.2 43.7% 48.2 45.6% 30.5 28.90/0 1.04 17 48 2.8 79.0 30.0 38.0% 31.5 39.90/0 26.5 33.5% 1.05 Torrens 1987, PU31711 [M] 98.9 38.0 38.4% 38.0 38.4% 36.0 36.4% 1.00 20 39 2.0 pI. 8, fig. 2 75.9 27.4 36.1% 29.6 39.0% 29.2 38.5% 1.08 24 42 1.8 Sturani 1967, PU31719 [M] 93.3 42.8 45.90/0 39.4 42.2% 28.6 30.7% 0.92 21 46 2.2 pI. 15, fig. 1 77.9 31.8 40.8% 31.6 40.6% 24.6 31.6% 0.99 19 44 2.3 PU31720 Figs. 12A-B [M] 67.4 23.5 34.90/0 25.9 38.4% 28.7 42.6% 1.10 22 36 1.6 52.0 18.1 34.8% 20.6 39.6% 22.9 44.0% 1.14 20 34 1.7 PU31680 Figs. 10G-H [m] 61.0 21.5 35.2% 21.8 35.7% 25.0 41 .0% 1.01 20 PU31677 Figs. 101-J [M?] 55.0 20.1 36.5% 23.5 42.7% 23.0 41 .8% 1.17 26 PU31681 Figs. 10E-F [m] 52.0 18.0 34.6% 19.5 37.5% 22.8 43.8% 1.08 21 36 1.7 38.0 13.0 34.2% 14.2 37.4% 18.1 47.6% 1.09 20 31 1.6 Figs. 10A-D PU31676 [m] 49.0 15.6 31.8% 20.5 41.8% 21.2 43.3% 1.31 24 hoiotype PU31682 Figs. 1 OK-M [m] 44.0 15.0 34.1% 16.7 38.0% 20.5 46.6% 1.11 20 33 1.7 33.0 10.0 30.3% 12.5 37.90/0 16.1 48.8% 1.25 23 34 1.5 Pavia et al. 2008, PU111577 [M] 39.2 13.0 33.2% 17.0 43.4% 18.0 45.90/0 1.31 22 pI. 3, figs. 9-10 D. Measurements for the specimens of Protozigzagiceras tethycum sp. novo Specimens Figures D H HID W WID U UiD Will Nil2 Nel2 PU31694 Figs. 16H-I [M] 51.0 17.0 33.3% 17.0 33.3% 20.1 39.4% 1.00 20 39 2.0 37.0 14.0 37.8% 14.0 37.8% 15.9 43.0% 1.00 21 PU1 11299 Figs. 16E-G [m] 40.0 14.6 36.5% 11.6 29.0% 17.1 42.8% 0.79 23 46 2.0 28.0 9.0 32.1 % 8.1 28.90/0 12.9 46.1% 0.90 19 Figs. 16J-L PU1 11574 [m] 36.0 10.0 27.8% 12.4 34.4% 17.4 48.3% 1.24 19 33 1.7 hoiotype PU31695 Figs. 16A-D [m] 30.0 9.7 32.3% 10.5 35.0% 13.6 45.3% 1.08 19 36 1.9 33.0 10.0 30.3% 12.5 37.90/0 16.1 48.8% 1.25 23 34 1.5 E. Measurements for the specimens of Protozigzagiceras fle xum sp. novo Specimens Figures D H HID W WID U UID Will Ni/2 Nei2 PU112319 Figs. 17E-F [M] 51.3 16.3 31.8% 14.9 29.0% 21.6 42.1% 0.91 41.5 13.7 33.0% 14.1 34.0% 17.4 41.9% 1.03 24 46 1.9 Figs. 17A-D PU111326 [m] 41.0 13.1 32.0% 15.0 36.6% 19.0 46.3% 1.15 hoiotype 30.0 9.2 30.7% 11.0 36.7% 14.6 48.7% 1.20 24 38 1.6 F. Measurements for the sEecirnens of Protozis.zas"iceras densum sE. nov o Specimens Figures D H HID W WID U UID WIH Nil2 Nel2 Figs. 18A-D PU111322 [m] 35.6 11.5 32.3% 13.8 38.8%) 16.1 45.2% 1.20 18 46 2.6 hoiotype G. Measurements for the sEecimens of Mes.a zis.z as.i ceras subarabicum sE. novo Specimen Figures D H HID W WID U U/D Will Nil2 Nel2 Figs. 19A-B FSL 177 639 [m] 100.0 22.3 22.3% 25.0 25.0% 60.0 60.0% 1.12 16 hoiotype ent 10 mm.