(Cretaceous) Ammonite Pachydesmoceras Maroccanum from North America

N. Jb. Geol. Paläont. Abh. 283/1 (2017), 77–84 Article E Stuttgart, January 2017

First record of the Cenomanian (Cretaceous) ammonite Pachydesmoceras maroccanum from North America

Paul L. Sealey and Spencer G. Lucas With 2 figures and 2 tables

Abstract: A single specimen of Pachydesmoceras maroccanum Collignon was recovered from the middle Cenomanian Acanthoceras amphibolum Zone in the Paguate Member of the Dakota Forma- tion in Sandoval County, New Mexico. Previous to this report, P. maroccanum was only known from the Cenomanian of the Tarfaya Basin in Morocco. The New Mexican specimen is the first known occurrence of P. maroccanum in North America, with only a few occurrences of Pachydesmoceras reported from the continent. It is also the first Cenomanian record of the genus from North America and indicates a trans-Atlantic migration of P. maroccanum.

Key words: Pachydesmoceras maroccanum, Dakota Formation, Paguate Member, Cenomanian, North America, USA.

1. Introduction Mexico. Thus, it is the first Cenomanian record of the genus from North America and the first occurrence of The Cretaceous ammonoid genus Pachydesmoceras P. maroccanum known from the continent. Here, we has not been previously recorded from the Cenomanian document this species in New Mexico and compare it of North America. In the USA, species of Pachydes- to other species of the genus. moceras have only been reported from the upper Al- bian and the upper Turonian (Cobban & Kennedy 1991: Abbreviations and conventions: Institution housing fossil collections used in this report: NMMNH refers to the New A2). P. colusaense Anderson from the probable upper Mexico Museum of Natural History and Science, Albuquer- Albian of California is the oldest known member of que. All dimensions are given in millimeters; D = diameter, the genus from North America. P. linderi (de Gros- U = umbilical diameter, Wh = whorl height, Wb = whorl souvre) from the upper Turonian of Colorado is the breadth, c = costal and ic = intercostal. youngest known Pachydesmoceras from the continent. P. maroccanum Collignon was originally described from the middle to upper Cenomanian of the Tarfaya 2. Locality and stratigraphy Basin in Morocco (Collignon 1966: 26, 58-59, pl. 12, fig. 4). A specimen of P. maroccanum was recently The San Juan Basin of northwestern New Mexico ex- recovered from the middle Cenomanian Acanthoceras poses a thick and very fossiliferous succession of Upper amphibolum Zone near the top of the Paguate Mem- Cretaceous marine, marginal marine and nonmarine ber of the Dakota Formation in Sandoval County, New strata that range in age from middle Cenomanian to

eschweizerbart_xxx 78 P.L. Sealey and S.G. Lucas late Maastrichtian. The basal portion of this section are strata of the intertongued Dakota-Mancos succession, which record the initial transgression of the Western Interior seaway into northwestern New Mexico (e.g., Lucas et al. 1998). The most precise age calibration of the Dakota-Mancos strata comes from ammonoid biostratigraphy (e.g., Cobban 1977a, b). The ammonoids indicate the entire succession is of middle-late Cenomanian age, encompassing the ammonoid zones of Conlinoceras tarrantense through Calycoceras canitaurinum (Table 1; e.g., Cobban 1977a, b; Sealey & Lucas 2016). This long and well known ammonoid succession has produced few new or rare taxonomic records during the last 40 years, so the rare species of ammonoid reported here is an unusual and singular occurrence. In northwestern New Mexico, the base of the Cre- taceous section is the intertongued Dakota-Mancos succession that consists of sandstone-dominated members of the Dakota Formation interbedded with shale-dominated members of the Mancos Formation. The succession, in ascending order, is Oak Canyon and Cubero members (Dakota), Clay Mesa Member (Man- cos), Paguate Member (Dakota), Whitewater Arroyo Member (Mancos) and Twowells Member (Dakota), and these strata contain ammonoids of middle-late Cenomanian age (e.g., Dane 1960; Landis et al. 1973; Cobban 1977a, b; Lucas et al. 1998; Sealey & Lucas 2003). The specimen of Pachydesmoceras maroccanum documented here came from NMMNH locality 9575 in the southeastern San Juan Basin of northwestern New Mexico, near the village of San Ysidro in San- doval County. This locality is in the upper third of the Paguate Member of the Dakota Formation, stratigraphi- cally above the Clay Mesa Member of the Mancos Formation, which yields ammonoids (notably Acan- thoceras amphibolum) and other marine invertebrate fossils from two stratigraphic levels (Fig. 1). Elsewhere, the Paguate Member also yields ammonoids of the A. Fig. 1. Location map of New Mexico and measured strati- amphibolum Zone (e.g., Cobban 1977a, b; Sealey & graphic section at the locality of Pachydesmoceras maroccanum. The section displays part of the intertongued Dakota- Lucas 2003). The A. amphibolum Zone is of middle Mancos succession of northwestern New Mexico, the Cubero Cenomanian age (Table 1; Cobban et al. 2006, fig. 1; and Paguate members of the Dakota Formation interbedded 2008, fig. 2; Merewether et al. 2011, fig. 2). with the Clay Mesa Member of the Mancos Formation. At NMMNH locality 9575, the stratigraphic section (Fig. 1) begins at the top of the Cubero Member of the Dakota Formation, tabular bedded and bioturbated, meter or more in diameter in its lowermost and upper fine- to medium-grained quartz sandstone. The overly- strata. A calcareous sandstone bed about 5 m above the ing Clay Mesa Member of the Mancos Formation is base of the Clay Mesa Member yields the ammonoid ~ 18 m thick and is mostly gray, slightly sandy shale species Acanthoceras amphibolum, indicative of the with intervals of limestone septarian nodules up to a eponymous middle Cenomanian ammonoid zone. Sep-

eschweizerbart_xxx First record of the Cenomanian (Cretaceous) ammonite Pachydesmoceras maroccanum 79 tarian nodules near the top of the Clay Mesa Member as in Pachydesmoceras (Matsumoto 1988: 22; Wright et also yield fossils of A. amphibolum. al. 1996: 75). The Paguate Member at NMMNH locality 9575 Faint to weak constrictions are present on the adapical side of primary ribs on the earlier part of the outer whorl, is about 9 m thick (its top is an erosionally stripped thus forming collared ribs. Differentiation of ribs into pri- surface). The entire unit is fine- to medium-grained maries and secondaries does not begin until approximately quartz sandstone that is ripple laminated or wavy bed- after the first one-sixth of the outer whorl. Strong, distant, ded except near the top of the unit, where the sandstone slightly sinuous rectiradiate primary ribs start at the sharply is laminar. NMMNH locality 9575 is in a limestone rounded umbilical shoulder, cross flattened flanks on the earlier part of the outer whorl and slightly convex flanks septarian nodule horizon about 2.5 m below the local on the later part of the outer whorl, where they are strong, top of the Paguate Member. bar-like and high across the venter. The primaries are distant on the earlier part of the whorl and more closely spaced on the later part of the outer whorl, for a total of 13 primaries per whorl. Increase in primaries and decrease in intercala- 3. Systematic paleontology tories in an adoral direction is gradual and fairly consistent. Some intercalatories start outside the umbilical shoulder, but most start on the outside third of the flank. A few secondar- Order Ammonoidea Zittel, 1884 ies, mostly on the earlier part of the outer whorl, appear to Suborder Ammonitina Hyatt, 1889 split on the outer flank, but it is difficult to make a definite determination. There are a total of 60-61 intercalated ribs. Superfamily Desmoceratoidea Zittel, 1895 On most of the phragmocone intercalated ribs crossing the Family Desmoceratidae Zittel, 1895 venter are weaker than the primaries but become stronger Subfamily Puzosiinae Spath, 1922 over the venter and are almost as strong as the primaries on Genus Pachydesmoceras Spath, 1922 the last preserved portion of the outer whorl. Ribs project only slightly forward over the venter and flare on the ven- Type species: Ammonites denisonianus Stoliczka, 1865, ter on the last half whorl. The suture is poorly preserved, p. 133, pl. 65, fig. 4; pl. 66; pl. 66a, by original designation but, where visible, is comparable to Puzosia in having long, (Spath 1922: 127). highly dissected saddles and lobes (e.g., Wright & Kennedy 1984, text-figs. 2h, m, 4b).

Pachydesmoceras maroccanum Collignon, 1966 Fig. 2; Table 2 4. Discussion and conclusions

1966 Pachydesmoceras maroccanum Collignon, p. 26, pl. Members of the subfamily Puzosiinae Spath, 1922 are 12, fig. 4. uncommon in the Upper Cretaceous of the Western 1987 Pachydesmoceras maroccanum Collignon. – Zabor- Interior and Gulf Coast (Cobban & Kennedy 1991: A1). ski, p. 36. 1991 Pachydesmoceras maroccanum Collignon. – Cob- The only other Cenomanian member of the subfam- ban & Kennedy, p. A2. ily from these regions is one unillustrated fragment of Puzosia (Puzosia) sp. from the middle Cenoma- Material: NMMNH P-71327 from locality 9575 in the nian Acanthoceras bellense Zone in Texas (Kennedy middle Cenomanian Acanthoceras amphibolum Zone in the & Cobban 1990: 90; Cobban & Kennedy 1991: A1). Paguate Member of the Dakota Formation, Sandoval County, New Mexico. The genus Pachydesmoceras ranges from the Albian to the Santonian and has been recorded from Poland Description: The specimen (Fig. 2) is a moderately-pre- (Kin & Niedźwiedzki 2012), Spain, France, Switzer- served mold of Pachydesmoceras maroccanum that ap- land, Germany, Yugoslavia, Romania, Morocco, Zu- pears to be all phragmocone, but because of preservation and luland, Madagascar, Iran, southern India, Japan, New weathering it is difficult to make a definite determination. It is fairly evolute with a moderately wide umbilicus and a Zealand, California and Colorado (Cobban & Kennedy low, vertical umbilical wall. Less than one-half of the previ- 1991: A1). ous whorl is covered by the next whorl. Inner whorls are not NMMNH P-71327 is comparable to Pachydesmo- exposed. The shell is compressed and high whorled on the ceras maroccanum Collignon, 1966 from the Ceno- outer whorl, with convergent flanks. The greatest breadth is manian of the Tarfaya Basin in Morocco in the slightly below mid-flank. It has a fairly narrowly arched and rounded flexuous, coarse, rectiradiate primary ribs that start at venter with the siphuncle exposed in many places. The shell has a costal diameter of 194 mm (Table 2). It has two distinct the umbilical shoulder, in having some intercalatories series of long and short ribs that project slightly forward that start near the umbilical shoulder and others that over the outer part of the whorl and that coarsen gradually, begin higher on the flank (Collignon 1966: 26). There

eschweizerbart_xxx 80 P.L. Sealey and S.G. Lucas

Fig. 2. Pachydesmoceras maroccanum from the Paguate Member of the Dakota Formation in Sandoval County, New Mexico, A, ventral, B, lateral and C, anterior views, NMMNH P-71327 from locality 9575. Middle Cenomanian Acanthoceras amphibolum Zone. Scale equals 2 cm.

Table 1. Zonal table showing some of the ammonite zones mentioned in the text that are present in the study area (modified from Cobban et al. 2008, fig. 2; Merewether et al. 2011, fig. 2).

Stage Substage Ammonite Zone Formation/Member Upper Calycoceras canitaurinum Dakota Formation/Two Wells Member Plesiacanthoceras wyomingense Dakota Formation/Paguate Member Acanthoceras amphibolum Mancos Formation/Clay Mesa Member Middle Acanthoceras bellense Plesiacanthoceras muldoonense Acanthoceras granerosense Cenomanian (pars) Dakota Formation/Cubero Member (lower)/ Oak Canyon Conlinoceras tarrantense Member are also no very clear bifurcated intercalary ribs, as in 26). Also, the holotype of P. maroccanum has 7 to 9 P. maroccanum (Collignon 1966: 26). The New Mexi- intercalated ribs between primaries (Collignon 1966: can specimen also resembles P. maroccanum in hav- 26), whereas the NMMNH specimen has 2 to 9 inter- ing slightly convex flanks and vertical umbilical walls calatories. However, the New Mexico specimen is much (Collignon 1966: 26). P-71327 seems to differ from the larger than the Moroccan one, so the variability could holotype of P. maroccanum in the outer whorl being be due to changes in growth. Consequently, the extent compressed instead of equal height and width and in of intraspecific variation in P. maroccanum cannot be being a little more evolute (Table 2; Collignon 1966: determined because of the very limited sample.

eschweizerbart_xxx First record of the Cenomanian (Cretaceous) ammonite Pachydesmoceras maroccanum 81

1 2 3 4 Table 2. Measurement chart of species of Pachydesmoceras. Collignon 1966; Collignon 1961; Matsumoto 1988; Col- 5 lignon 1964; Anderson 1958; NMMNH refers to the New Mexico Museum of Natural History and Science, Albuquerque. Measurements are in millimeters. Numbers in parentheses are percentages of diameter. D = diameter, U = umbilical diameter, Wh = whorl height, Wb = whorl breadth and H/h = measurements of whorl height indicating rate of whorl expansion in 180 degrees, e = estimated, c = costal and ic = intercostal.

Taxon Specimen D Wb Wh Wb:Wh U H/h NMMNH P-71327 Pachydesmoceras maroccanum 194.0 68.8 (35.5) (e) 81.5 (42.0) 0.84 66.6 (34.3) – (c) “ “ (ic) 192.1 63.8 (33.2) (e) 78.3 (40.8) 0.81 66.6 (34.7) 1.47 “ Holotype(1) 109 45 (41.3) 45 (41.3) 1.00 32 (29.0) – Pachydesmoceras rarecostatum Holotype(2) 163 72 (44.1) 69 (42.3) 1.04 53 (32.5) – Pachydesmoceras hourcqi Holotype(2) 215 84 (39.1) 92 (42.8) 0.91 66 (30.7) – Pachydesmoceras pachydiscoide Holotype(3) 215 83 (38.6) 98 (45.6) 0.85 65 (30.2) 1.75 Collignon, 1961, Pachydesmoceras denisonianum (3) 166 65 (39.1) 67 (40.4) 0.97 52 (31.3) 1.43 pl. 8, fig. 1, 1a Pachydesmoceras? hottingeri Holotype(1) 187 68 (36.4) 84 (45.0) 0.81 50 (26.7) – Pachydesmoceras kossmati Holotype(3) 168 67 (39.9) 71 (42.3) 0.94 47 (28.0) 1.42 Collignon, Pachydesmoceras linderi (1) 114 45 (39.5) 46 (40.3) 0.98 35 (30.7) – pl. 18, fig. 4

(4) 111 (47.2) (c) 0.97 (c) Pachydesmoceras radaodyi Holotype 235 108 (45.9) 62 (26.4) – 102 (43.4) (ic) 1.06 (ic) Pachydesmoceras mihoense Holotype(3) 145 39 (26.9) 48.4 (33.4) 0.81 59 (40.7) 1.29 Pachydesmoceras colusaense Holotype(5) 239 83.3 (34.8) 106.5 (44.6) 0.78 76 (31.8) –

NMMNH P-71327 differs from Pachydesmoceras and slightly convex flanks on the outer whorl instead linderi (de Grossouvre, 1894) from the Turonian and of inflated flanks and in the stronger bar-like primaries Coniacian of France and upper Turonian of Madagas- being much stronger than the secondaries (Matsumoto car, Morocco, Romania and Colorado in having rectira- 1988: 22, 119, figs. 50-54); from P. rarecostatum Colli- diate ribs of varying strength and more intercalatories gnon, 1961 from the Cenomanian of Madagascar in be- (see Collignon 1966, pl. 18, fig. 4; Cobban & Kennedy ing compressed instead of depressed (Collignon 1961: 1991: A2; Robaszynski et al. 2014: 132; Kennedy et al. 40, pl. 9, fig. 1); and from P. hottingeri Collignon, 1966 2015: 454); from P. denisonianum (Stoliczka, 1865) from the Albian of Morocco in the coarser ribbing, less from the upper Albian and Cenomanian of southern intercalatories and less branching on the outer flanks India and Japan, upper Cenomanian of North Germany, (Collignon 1966: 26, pl. 2, fig. 1). Cobban & Kennedy lower Cenomanian and lower Turonian of Madagas- (1991: A2) state that P. hottingeri belongs to some other car and lower Turonian of Nigeria and Spain in ribs genus of Puzosiinae. having a weaker ventral projection, the primaries be- Pachydesmoceras hourcqui Collignon, 1961 from ing more distant adapically and more closely spaced the upper Turonian of Madagascar also has 13 prima- adorally and in having an increase in primaries and ries on the outer whorl but differs in these ribs being decrease in secondaries in an adoral direction (see Col- weaker and in only having a maximum of 5 intercala- lignon 1961, pl. 8; Szász 1986: 121; Zaborski 1987: 34; tories (Collignon 1961: 42, pl. 11). P. pachydiscoide Matsumoto 1988: 109, figs. 48-49; Kaplan et al. 1998: Matsumoto, 1954 from the Turonian of Sakhalin, Ja- 84; Küchler 1998: 165, 180, 183, 192, 194, 198, 203); pan, California, southern India and Madagascar and from P. kossmati Matsumoto, 1987 from the lower and lower Coniacian of Madagascar and northern KwaZu- middle Cenomanian of Madagascar (Kennedy et al. lu-Natal does not have the strong, bar-like primaries 2013: 632) and lower Turonian of southern India, Japan of the NMMNH specimen, and it has long ribs that and Sergipe, Brazil in the less sinuous ribs (Gale et al. are more distantly spaced at a similar diameter, ribs 2005: 172), in the flattened flanks on the inner whorl that have a stronger ventral projection and constric-

eschweizerbart_xxx 82 P.L. Sealey and S.G. Lucas

tions to a greater diameter (Matsumoto 1954: 102, pl. America and indicates a trans-Atlantic migration of P. 9, fig. 2; Matsumoto 1988: 132, figs. 56-60; Kennedy maroccanum. & Klinger 2014: 17). P. colusaense Anderson, 1902 from the likely upper Albian of California does not have strong bar-like primaries, has more flexuous ribs Acknowledgements and is more involute (Anderson 1902: 96-97, pl. 5, figs. 128-129, pl. 10, fig. 200; Anderson 1958: 236, pl. 10, We thank Michael Foley, who discovered and donated the fig. 1). P. mihoense (Matsumoto, 1954) from the up- Pachydesmoceras to the NMMNH and took the authors to per Coniacian-lower Santonian of Sakhalin and Japan the locality. Comments by Christina Ifrim and an anony- mous reviewer on an earlier version of this manuscript im- and Coniacian of Russia has strong, very coarse, thick, proved its content. distant, flared ribs (Matsumoto 1988: 137, figs. 61-65; Zonova & Yazykova 1998: 490-491, fig. 5). P. radaodyi Collignon, 1964 from the middle Cenomanian of Madagascar has distant, flared primary ribs that are References ol extraordinarily strong on the adult body chamber (C - Anderson, F.M. (1902): Cretaceous deposits of the Pacific lignon 1964: 58, pl. 333, fig. 1498; Matsumoto 1988: coast. – California Academy of Science Proceedings, 3rd 23, 141; Cobban & Kennedy 1991: A2). See Table 2 series, Geology, 2 (1): 1-131. for a measurement chart of comparable species of Anderson, F.M. (1958): Upper Cretaceous of the Pacific Pachydesmoceras. coast. – Memoirs of the Geological Society of America, 71: 378 pp. ollignon C (1966: 58-59) listed several ammonites Brongniart, A. (1822): Sur quelques terrains de Craie hors from the same deposit in the Tarfaya Basin in Morocco du Bassin de Paris. – In: Cuvier, G. & Brongniart, A. that were presumably associated with Pachydesmoc- (Eds.): Description géologique des environs de Paris (3rd eras maroccanum. 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eschweizerbart_xxx First record of the Cenomanian (Cretaceous) ammonite Pachydesmoceras maroccanum 83

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(2012): First record of the puzosi- lopoda – Palaeontographical Society Monographs, 68: ine ammonite genus Pachydesmoceras from the middle 1-26 (1853), 11-16 (1855), 17-27 (1857). and upper Turonian of Poland. – Cretaceous Research, Spath, L.F. (1922): On the Senonian ammonite fauna of Pon- 33: 15-20. doland. – Transactions of the Royal Society of South Af- Kossmat, F. (1895-1898): Untersuchungen über die südindi- rica, 10: 113-147. sche Kreideformation – Beiträge zur Paläontologie und Stoliczka, F. (1864-1866): The fossil Cephalopoda of the Geologie Österreich-Ungarns und des Orients, 9 (1895): Cretaceous rocks of southern India (Ammonitidae). – 97-203 (1-107), pls. 15-25 (1-11); 11 (1897): 1-46 (108- Memoirs of the Geological Survey of India, 1: 41-216. 153), pls. 1-8 (12-19); 12 (1898): 89-152 (154-217), pls. Szász, L. (1986): Lower Turonian ammonite assemblage 14-19 (20-25). in the Maramureş Mountains (East Carpathians – Ro- Küchler, T. (1998): Upper Cretaceous of the Barranca (Na- mania). – Institutul de Geologie şi Geofizică, 70/71 (3): varra, northern Spain); integrated litho-, bio- and event 117-134.

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Wright, C.W. & Kennedy, W.J. (1984): The Ammonoidea of Zonova, T.D. & Yazykova, E.A. (1998): Biostratigraphy and the Lower Chalk, Part I. – Monographs of the Palaeon- correlation of the Turonian-Coniacian succession and the tographical Society: 1-126. Turonian-Coniacian boundary problem in the Far East Wright, C.W. & Kennedy, W.J. (1987): The Ammonoidea of Russia based on ammonites and inoceramids. – Acta the Lower Chalk, Part 2. – Monographs of the Palaeon- Geologica Polonica, 48: 483-494. tographical Society: 127-218. Wright, C.W., Callomon, J.H. & Howarth, M.K. (1996): Manuscript received: June 17th, 2016. Cretaceous Ammonoidea. – In: Kaesler, R.L. (Ed.): Revised version accepted by the Bremen editor: December Treatise on Invertebrate Paleontology, Part L, Mollusca 11th, 2016. 4, Cretaceous Ammonoidea. – 362 pp.; Boulder & Law- rence (Geological Society of America & University of Address of the authors: Kansas Press). Zaborski, P.M.P. (1987): Lower Turonian (Cretaceous) am- Paul L. Sealey, Spencer G. Lucas, New Mexico Museum monites from south-east Nigeria. – Bulletins of the Brit- of Natural History and Science, 1801 Mountain Road, NW, ish Museum of Natural History, (Geology) 41 (2): 31-66. Albuquerque, NM 87104, USA; Zittel, K.A. (1884): Cephalopoda. – In: Handbuch der Pa- e-mails: [email protected]; laeontologie, Abteilung 1, 2 (3): 329-522; München & [email protected] Leipzig (Oldenbourg). Zittel, K.A. (1895): Grundzüge der Palaeontologie (Palaeo- zoologie). – vii + 972 pp.; München & Leipzig (Olden- bourg).

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