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Species Anapachydiscus fresvillensis Seunes 1890



Diagnosis / Definition:
Kennedy (1986):
Description. All specimens studied are internal moulds ofmacroconeh phragmoeones, 67-148 mm in diameter. Coiling involute, umbilieus eomprising 19•8-22•4% of diameter with 60% of previous whorl covered. Whorl section depressed, reniform in juvenile (breadth to height ratio up to 1-12) with greatest breadth at umbilical shoulder, umbilical wall rounded and undercut, inner flank rounded, outer converging to broadly rounded venter. Whorl breadth to height ratio decreases through ontogeny and section is slightly compressed from 120-130 mm diameter. Up to 60-65 mm (PI. 9, figs. 1-3), ten or eleven low, broad ribs arise at umbilical seam and develop into prominent umbilical bullae. These give rise to paired (rarely three) ribs, while occasional non-tuberculate ribs arise at shoulder and shorter intercalatories arise on inner flank to give a total of thirty-one or thirty-two ribs per whorI. These are narrow and prorsiradiate on inner flank at smatlest diameter visible, curve forwards over ventrolateral shoulder and cross venter in broad convexity, attenuating and effacing over mid¬venter. As size increases (pI. 8) ribs strengthen and are strongest over venter by a diameter of 50 mm. Bullae decline beyond 60-65 mm and migrate out to inner flank position (PI. 8, fig. 2), giving rise to pairs of ribs with, in addition, both long and short intercalatories that strengthen markedly over venter; there are forty ribs at 115 mm. Umbilical bullae decline beyond 120 mm and ribs progressively efface on inner and middle flank so that, by largest diameter seen in the present material (the lectotype; PI. 7), ornament is confined to coarse, broadly convex, rounded ribs, forty per whorl, on ventrolateral shoulders and venter only, with smooth flanks on mould at least. The suture (text-fig, 4A) is finely subdivided.
Ward & Kennedy (1993):
Description. - All specimens crushed, and original whorl section cannot be reconstructed. Coiling very involute with small umbilicus comprising around 20 percent of diameter. Whorls high, expanding rapidly; in juveniles to diameter of 60-70 mm, estimated 10-1 1 ribs per whorl arise at umbilical seam and strengthen into large conical tubercles perched on umbilical shoulder. These are flat topped and were septate bases of umbilical spines, occasionally visible on external molds in field. Tubercles give rise to one or two narrow, sharp, distant primary ribs, straight and prorsiradiate on inner to mid-flank, where occasional short ribs intercalate, to give a total of 30-32 ribs per whorl. Ribs flex forward on ventrolateral shoulder and strengthen, crossing venter in broad convexity. As size increases, umbilical tubercles weaken and migrate out to inner flank position. They give rise to pairs of ribs, with both long and short intercalatories that are weak on mid-flank but strengthen markedly on outer flank and venter where they are narrow, rounded, and separated by wide interspaces, totalling up to 23 per half whorl. Beyond 120-140 mm diameter, bullae decline, as does inner flank ornament, so that large specimens have coarse distant concave ribs on ventrolateral shoulders that cross venter in broad convexity. Adult specimens up to 300 mm in diameter, and may have up to 50 ventral ribs.
Discussion / Comments:
Kennedy (1986):
Discussion. Massive whorls, depressed and with strong umbilical bullae giving rise to groups of ribs, plus the effacement of ornament on all but the ventrolateral shoulders and venter in rniddle growth readily distinguish macroconch A. [resvillensis from the other pachydiscids present in the Ca1caire a Baculites. Most workers refer the species to Pachydiscus, but the massive, involute whorls and the presence of umbilical spines and tubercules giving rise to paired ribs suggest it is better placed in Anapachydiscus. Of the European species, the present form has generally been confused with P. colligatus (Binkhorst, 1861). This species is discussed at length elsewhere in my revision of the ammonites of the type Maastrichtian (in prep.). Suffice it to say that the proposed lectotype (the original of Binkhorst 1861, pI. 8, which is also the holotype by monotypy of P. vandenbroecki de Grossouvre, 1894, p. 207) is a much less massive shell, more evolute, and slower expanding; moreover, on moulds, the primary ribs extend to the umbilical shoulder and the secondaries extend weIl down the flank at a diameter where they have disappeared in P. (P.) [resvillensis, while the ventral ribbing is much more subdued in middle growth. The smallest topotype of P. (P.) colligatus seen is over 200 mm in diameter, so that comparison ofthe early stages is difficult, The reader is referred to the revision ofthe Maastricht fauna noted above for additional discussion. P. quiriquinae Steimnann, 1895 (p. 74, pI. 6, fig. 3; text-fig. 5) from the Upper Maastrichtian of Quiriquina Island, Chile (lectotype, here designated, the original ofSteimnann 1895, pI. 6, fig. 3) has the following dimensions (after Steinmann): The style of ornament is identical to that of A. fresvillensis, and there are, according to Steimnann, eleven umbilical tubercles and forty to forty-five ventral ribs as opposed to thirteen to fifteen umbilical tubercIes and forty-eight ventral ribs in [resvillensis. An exarnination of the lectotype of fresvillensis shows identical ornament in both form and density, although the lectotype of quiriquinae is more compressed (Wb: Wh ratio is 0.77 vs. 0,94). I regard them as conspecific. P. supremus Pethö, 1906 (p. 88, pl. 5, fig. I) is also a macroconch fresvillensis; it is from the Maastrichtian ofFruska Gora, Yugoslavia. The sketchily figured Brazilian P. sumneri, P. poseidon, and Canadoceras riogramense of Maury (1930) may belong here, but they are scarcely recognizable from the figures; they are Maastrichtianin age. P. subrobustus (Seunes, 1891) (p. 15, pl. 13 (2), fig. 1) is immediately distinguished by its evolute coiling, coarse ribs, and lack of umbilical bullae/spines. A. wittekindi(Schlüter, 1872) (see Blaszkiewicz 1980, p. 50, pl, 42, figs. 1 and 2; pI. 43, fig. 2; pls, 44-47; pI. 48, figs. 3 and 4; pI. 49, figs. 1 and 3; pI. 50, figs. 2 and 3; pls. 51-53) and A. vistulensis Blaszkiewicz, 1980 (p. 48, pI. 42, figs. 3 and 4; pI. 43, figs. land 3; pI. 48, figs. 1 and 2) both have much more coarsely ribbed nuclei than the present form, while P. wittekindi has a middle feebly ribbed and an adult strongly ribbed growth stage and P. vistulensis a strongly ribbed adult stage, all features which immediately distinguish them from the present species. • P. auritocostatus of Seunes (1890b, p. 239, pI. 8, fig. 4) non Schlüter, a diminutive bituberculate form that occurs with A. fresvillensis at the Carriere des Bernes between Gan and Rebenacq in the Pyrenees-Atlantiques, may be the rnicroconch of this species; I have, however, failed to locate the original specimen and can do no more than suggest the possibility
Ward & Kennedy (1993):
Type. - Lectotype, designated by-Kennedy, 1986c, p. 44, is No. A1 186 in the collections of the Ecole des Mines, Paris, now in the collections of the Université Claude-Bernard, Lyon, the original of Seunes, 1980a, p. 3, Pl. 2(1), fig. 1, from the Calcaire 21 Baculites of Fresville, Manche, France (Figure 36.1, 36.2). Discussion. - The lectotype is an internal mold (Figure 36.1, 36.2), and ornament is relatively subdued. External molds from the Maastricht area (Kennedy, 1987, Pl. 10, figs. 1-3, Pl. 23, fig. 5) show the strong umbilical spines that indicate fresvillensis to be an Anapachydiscus, as well as sharp distant ribs like those seen on the composite molds from the Biscay sections. Stinnesbeck (1986, p. 221, Pl. 14, fig. 1, Pl. 16, fig. 1, text-fig. 28b) has recently redescribed material from central Chile as Anapachydiscus fresvillenis quiriquinae (Steinmann, 1895). For reasons outlined by Kennedy (1986c, 1987), we regard this as a synonym of A. fresvillensis sensu stricto. Through the kindness of Professor A. A. Atabekian (Leningrad) we have been able to study casts of species described by Atabekian and Akopian (1 969). Pachydiscus colligatus michailovi Atabekian and Akopian, 1969 (p. 14, Pl. 5, fig. 2, Pl. 6, fig. 2, Pl. 7, figs. 1, 2) is based on crushed composite molds of the present species, with the same distant ribbing, and the flat-topped bases of septate spines clearly visible in some specimens (Figure 37.1-37.6). A typical Armenian example referred to fresvillensis by these authors is shown in Figure 38.1-38.3. Differences between A.fresvillensis and European Campanian species of the genus are discussed by Kennedy (1 986c, 1987) and do not require repetition here. Sparse ribbing, umbilical spines, and massive shell form immediately distinguish the species from all other Pachydiscus described from the Biscay region. Anapachydiscus terminus n. sp., described below, has a comparable gross morphology, but is easily separated by the much finer, dense crowded ribs, up to 60 per whorl, that persist to up to 100 mm diameter, whereas fresvillensis has around 40-45 ribs at this size, the ribs already effaced on the middle of the flanks. Adults of both species show a restriction of ribs to ventrolateral shoulders and venter. Occurrence.- In the Biscay sections this taxon ranges from near the top of Member I (lower Maastrichtian) to low in Member IV (upper Maastrichtian). Where well dated elsewhere, the species is restricted to the upper Maastrichtian, with records from Tercis, Landes, inland sections in Pyrénés-Atlantiques, and Haute Garonne, the Calcaire à Baculites of the Cotentin Peninsula, Manche, France; Limburg, the Netherlands, Denmark, Yugoslavia, the Armenian SSR, southern India, Madagascar, western Australia, Chile, and possibly Brazil.
Systematics:

38
 Ordo Ammonoidea
  Subordo Ammonitina
   Superfamilia Desmocerataceae
    Familia Pachydiscidae
     Genus Anapachydiscus
      Species Anapachydiscus fresvillensis

39
  Ordo Ammonoidea
   Subordo Ammonitina
    Superfamilia Desmocerataceae
     Familia Pachydiscidae
      Genus Anapachydiscus
       Species Anapachydiscus fresvillensis
Synonym list:
Kennedy (1986):
1861 Ammonites colligatus Binkhorst. - Binkhorst : 25 (pars) pl. 6, fig. 3a-f(?); pl. 7, fig. 2c; pl. 8, figs. 1 and 2
1890 Pachydiscus fresvillensis Seunes. - Seunes : p.3 pl. 2(1), fig. 1
1890 Pachydiscus fresvillensis Seunes. - Seunes : p.236 pl. 7, fig. 1; pl. 8, figs. 1-3
? 1890 Pachydiscus auritocostatus Schlüter. - Seunes : p.239 pl. 8, fig. 4 (non Schlüter)
1891 Pachydiscus fresvillensis Seunes. - Seunes (non Seunes, 1890) : p.14 pl. 12 (3), fig. 1
1894 Pachydiscus colligatus Binkhorst. - Grossouvre : 202 (pars) pl. 24, figs. 1 and 3 only (non pl. 33, fig. 1)
1895 Pachydiscus quiriquinae Phillipi. - STEINMANN : p.74 pl. 6, fig. 3; text-fig. 5
1895 Pachydiscus fresvillensis Seunes. - STEINMANN : p.77
1906 Pachydiscus supremus Pethö. - Pethoe : p.88 pl. 5, fig. 1
1908 Pachydiscus colligatus Binkhorst. - Grossouvre : p.28 pl. 4, figs. 1-3; pl. 5, fig. 1; pl. 6, fig. 1 sp. emend. de Gross
? 1930 Pachydiscus sumneri Maury. - Maury : p.155 pl. 13, figs. 1 and 2
? 1930 Parapachydiscus poseidon Maury. - Maury : p.155 pl. 15
? 1930 Canadoceras riogramense Maury. - Maury : p.169 pl. 21, fig. 2
1930 Parapachydiscus sp. indet. Hyatt. - Wetzel : p.86 pl. 14, fig. 1
1938 Parapachydiscus fresvillensis Seunes. - COLLIGNON : 101 (51) pl. 7, figs. 4 and 5; text-figs. O and P
? 1952 Pachydiscus sp. aff. colligatus Binkhorst. - COLLIGNON : p.79 pl. 26, fig. 2
? 1955 Pachydiscus sp. aff. colligatus Binkhorst. - COLLIGNON : p.74 pl. 26, fig. 2
1969 Pachydiscus colligatus fresvillensis Seunes. - ATABEKIAN & AKOPIAN : p.13 pl. 6, fig. 1
1971 Pachydiscus fresvillensis Seunes. - COLLIGNON : p.30 pl. 652, fig. 2408
1985 Pachydiscus (Pachydiscus) fresvillensis Seunes. - Henderson & McNamara : p.78 pl. 8, figs. 3-6; pl. 9, figs. 1 and 2; text-figs. 12b, 13a, 15b
1986 Anapachydiscus fresvillensis Seunes. - Kennedy : p.42 pl. 7, 8, 9, figs. 1-3; text-figs. 3M,N,Q,4A
Ward & Kennedy (1993):
1890 Pachydiscus fresvillensis Seunes. - Seunes : p.3 pl. 2(1); fig. 1
1986 Anapachydiscus fresvillensis Seunes. - Kennedy : p.1008 pl. 1; fig. 4, 5 (with full synonymy)
1986 Anapachydiscus fresvillensis Seunes. - Kennedy : p.42 pl. 7; pl. 8; pl. 9, fig. 1-3; text-fig. 3M, N, Q, 4A
1986 Anapachydiscus fresvillensis Seunes. - Kennedy : p.160 fig. 8D, E
1986 Anapachydiscus fresvillensis quiriquinae Steinmann. - Stinnesbeck : p.221 pl. 14, fig. 1; pl. 16, fig. 1; text-fig. 28b
1987 Anapachydiscus fresvillensis Seunes. - Kennedy : p.173 pl. 6; pl. 7, fig. 1,2; pl.8; pl. 9, fig. 1, 2; pl. 10, fig. 1-5; pl. 11, fig. 5, 6; pl. 12, fig. 12-14; pl. 13, fig. 6, 7; pl. 14, fig. 1-3, 7, 11, 12; pl. 15, fig. 4-6; pl. 23, fig. 5.
1993 Anapachydiscus fresvillensis Seunes. - Ward & Kennedy : p. 41, 42, 43, 44, 4 fig. 35.3, 35.5, 35.6, 36.1, 36.2, 37.1-37.6, 38.1-38.3, 40.8, 45.1
Was used in synonym list of:
Anapachydiscus terminus
Menuites fresvillensis Seunes 1890
Stratigraphy - relative ages:
upper Maastrichtian - Maastrichtian: Kennedy (1986)
upper Maastrichtian - lower Maastrichtian: Ward & Kennedy (1993)
Specimen:
Collections of the Université Claude-Bernard - Lyon, Inventory number: No. A1186
References:

Binkhorst,J.T.. (1861):
Monographie des gastropodes et des céphalopodes de la Craie Supérieure du Limburg.
In: Maastricht and Muller Frères Eds: Muquardt, G.. p. 17,83,44

Seunes,J.. (1890):
Recherches géologiques sur les terrains secondaires et l' Eocene inferieur de la region sous-pyrénéene du sud-ouest de la France (Basses-Pyrenees et Landes). p. 250 pp.

Seunes,J.. (1890):
Contribution a l'etude des cephalopodes du Cretace Superieur de France. 1. Ammonites du Calcaire ä Baculites du Cotentin. . Mém. Soc. geol. Fr. Paleont. 1, Mém. Vol. 2 p. 1-7

Seunes,J.. (1891):
Contributions à l’étude des Céphalopodes du Crétacé Supérieur de France. Ammonites du Calcaire à Baculites du Cotenin (Suite); II. Ammonites du Campanien de la région sous-Pyrénéenne, Département de Landes . Mémoirs de la Société Géologique de France, Paléontologie Vol. 2(2) p. 8-22

Grossouvre,d.A.. (1894):
Recherehes sur la craie superieure (misdated 1893). .
In: Paleontologie. Les ammonites de la craie superieure, Mem. Servo Carte geol. det. Fr. Vol. 2 p. 264 pp.

STEINMANN,G.. (1895):
Die Cephalopoden der Quiriquina-Schichten.
In: Neues Jahrbuch für Mineralogie, Geologie undPaläontologie, 1895 p. 64-94

Pethoe,J.. (1906):
Die Kreide (Hypersenon-) fauna des Peterwardeiner (Pétervárader) Gebirges (Fruska Gora). . Palaeontographica Vol. 52 p. 57-336

Grossouvre,d.A.. (1908):
Description des ammonites du Cretace Superieur du Limbourg Belge et Hollandais et du Hainault . Mem. Mus. r. Hist. nat. Belg Vol. 4 p. 1-39

Maury,C.J.. (1930):
O Cretaceo da Parahyba do Norte. , Album das estampas p. 305

Wetzel,W.. (1930):
Die Quiriquina-Schichten als Sediment und paläontologisches Archiv . Palaeontographica Vol. 73 p. 49-104

COLLIGNON,M.. (1938):
Ammonites Campaniennes et Maastrichtiennes de I'ouest et du sud de Madagascar. . Annales Geologiques du Service des Mines de Madagascar Vol. 9 p. 55-118

COLLIGNON,M.. (1952):
Ammonites neocretacees du Menabe (Madagscar). II-Les Pachydiscidae. . Trav. Bur. geol. Madagascar Vol. 41 p. 144 pp.

COLLIGNON,M.. (1955):
Ammonites neocretacees du Menabe (Madagascar). II-Les Pachydiscidae. . Annls geol. Servo Mines Madagascar Vol. 21 p. 98 pp.

ATABEKIAN,A.A.. and AKOPIAN,v.T.. (1969):
Late Cretaceous ammonites of the Armenian SSR (Pachydiscidae). . Proceedings of the Armenian Academy of Science, Earth Sciences. [In Georgian]. Vol. 6 p. 3-20

COLLIGNON,M.. (1971):
Atlas des fossiles caracteristiques de Madagascar (Ammonites)(Maestrichtian). Vol. 17 p. 82

Henderson,R.A.. and McNamara,K.J.. (1985):
Maastrichtian non-heteromorph ammonites from the Miria Formation, Western Australia . Palaeontology Vol. 28 p. 35-88

Stinnesbeck,W.. (1986):
Zu den faunistischen und palökologischen Verhältnissen in der Quriquina Formation (Maastrichtium)Zentral-Chiles . Palaeontographica A194((4-6)) p. 99-237

Kennedy,W.J.. (1986):
Upper Campanian and Maastrichtian ammonites from the Petites-Pyrénées, southern France . Eclogae Geologicae Helvetiae Vol. 79 p. 1001-1037

Kennedy,J.W.. (1986):
The Campanian-Maastrichtian ammonite sequence in the environs of Maastricht (Limburg, the Netherlands) . Newsletters on Stratigraphy Vol. 16 p. 149-168

Kennedy,W.J.. (1986):
The ammonite fauna of the Calcaire à Baculites (Upper Maastrichtian) of the Cotentin Peninsula (Manche, France) . Palaeontology Vol. 29(1) p. 25-83

Kennedy,W.J.. (1987):
The ammonite faunas of the type Maastrichtian, with a revision of Ammonites colligatus Binkhorst, 1861 . Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre Vol. 56 p. 151-267

Ward,P.D.. and Kennedy,W.J.. (1993):
Maastrichtian Ammonites from the Biscay Region (France, Spain) . Memoir (The Paleontological Society), Journal of Paleontology34 (Supplement to Vol. 67)(5) p. 1-58

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