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Subspecies Hoploscaphites constrictus crassus Lopuski 1911 | ||||||
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| Diagnosis / Definition: | ||||||
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Machalski (2005): EMENDED DIAGNOSIS.- A temporal subspecies in the Hoplo− scaphites constrictus lineage, transitional between H. c. lviv− ensis subsp. nov. and H. c. johnjagti subsp. nov., defined by the absence of ribbing on a large sector of body chamber in adult macroconchs. |
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| Discussion / Comments: | ||||||
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Machalski (2005): TYPE MATERIAL.- A specimen from the upper upper Maastrichtian at Kazi− mierz Dolny, illustrated by Łopuski (1911: 115, pl. 2: 4, 5, pl. 3: 1, 2), presumably lost, was designated lectotype by Błaszkiewicz (1980: 37). MATERIAL.- From Poland: 88 specimens from Rejowiec (ZPAL Am. 12/92, 242, 364, 848–850, 853, 860–863, 866, 869, 875, 941, 943, 944, 947, 949, 950, 952, 954–963, 965–968, 970, 972–978, 980–984, 987, 989, 991–996, 998–1002, 1004–1010, 1012, 1015–1027, 1029–1032); 14 specimens from Podgórz (ZPAL Am. 12/246–248, 250, 251, 675, 676, 732, 735, 737, 739, 753, 757, 798); three specimens from Albrychtówka (ZPAL Am. 12/98 a, b, 728); 117 speci− mens from the town quarry south of Kazimierz Dolny (ZPAL Am. 12/30, 92–96, 101–105, 109–112, 115, 116, 119–121, 126, 127, 129–134, 137–147, 149–151, 153–156, 157–160, 624–632, 634–640, 643, 650–655, 657, 659–661, 664, 667–670, 677, 679, 681–685, 687, 689–691, 693, 694, 696–699, 725–727, 729–731, 746–751, 754, 810, 811, 813, 931, 1062); 123 from Nasiłów (ZPAL Am. 12/1–10, 12–16, 18–57, 59–63, 65, 67, 69, 71–77, 79, 82–87, 90, 91, 125, 161, 162, 308–315, 318, 320–322, 346, 363, 644–647, 806–809, 1217, 1218, 1220–1239); four specimens from Bochotnica (ZPAL Am. 12/311, 319, 324, 745); twenty−one from Kli− musin (ZPAL Am. 12/327–343, 352, 357–359); seven from Prawiedniki (ZPALAm. 12/350, 351, 353–355, 404, 424); six fromMętów (ZPAL Am. 12/344–349). From Denmark: three specimens from Bjerre (MGUH 1965.588, 1954.634, unregis− tered); 10 from Hov (MGUH 20156, 27743, 1965.568, 1965.569, 1965.575, 1965.577, 1965.578, 1965.585, 1973.399, 1973.400); 21 from Stevns Klint (MGUH 20206 and 20 un− registered specimens in the private collection of Alice Ras− mussen, Fakse). From the Ukraine: one specimen from Lviv (DPM NANU PZ−K−N9282); five from Griboviczi (formerly Grzybowice) (DPMNANUPZ−K−N9204, 9206, 9843–9845). DISCUSSION.- Łopuski (1911: 116) differentiated his Scaphi− tes constrictus Sowerby var. crassus from the nominal species only by the “enormous thickness of the conch”. The lectotype, an adultmacroconch, is a thick−set individual indeed (Łopuski 1911: 115, pl. 2: 4, 5, pl. 3: 1, 2), but its inflation is exaggerated by strong post−mortem deformation. According to Łopuski (1911), this specimen came from “Kazimierz”. Thus, it could have been collected from either the lower part of the Kazi− mierz Opoka as exposed at the town quarry south of Kazi− mierz Dolny or from the upper part of that unit, occasionally accessible during construction works in the town. Material from the Kazimierz Opoka studied comprises 118 adult macroconchs (Fig. 10B, F, G) and 33 adult micro− conchs (Fig. 7D, G, H). Two subsamples from relatively nar− row intervals have been measured; one is from a marly opoka, 2 metres thick, at the bottom of the upper third of the town quarry (referred to as the “Kazimierz Dolny sample” below), the other is from a hard limestone layer, up to one metre thick, which forms the top of the Kazimierz Opoka succession at Nasiłów (the “Nasiłów sample” below). The Kazimierz Dolny sample comprises 65 adult macro− and 15 adult microconchs; the former range between 29 and 66 mm, while the latter range between 22 and 38 mmin max− imum diameter (Fig. 9C). The Nasiłów sample includes 28 adult macro− and 8 microconchs, measuring between 38 and 73mmand between 24 and 35mmin maximum diameter, re− spectively (Fig. 9D). There is thus some overlap of size be− tween dimorphs of H. c. crassus in these samples (contrary to Makowski 1962, who claimed that there was no overlap in the material from the Kazimierz Dolny area). The material of H. c. crassus from the Kazimierz Dolny area shows a wide range of variation in conch proportions and ornament, which is best appreciated in illustrations sup− plied by Łopuski (1911), Błaszkiewicz (1980) and Machal− ski (1996, 2005). The development of ornament in adult macroconchs does not depend on the size of the conchs as the smaller specimens reveal essentially the same ornament as the larger individuals (Fig. 11A–C). In general, the Kazimierz Opoka material differs from that of H. c. constrictus from Cotentin in the ornament of the body chamber (Fig. 8). Flanks of the shaft are covered with ribs in most adult macroconchs from Cotentin (Kennedy 1986). In contrast, ribbing is absent from a large sector of the body chamber, including the shaft and a considerable region of the hook, in almost all adult macroconchs from the Kazi− mierz Opoka. Only faint striae or growth lines are visible on the flanks in better−preserved specimens, including those preserved as xenomorphic imprints and replicas on oyster shells (Lehmann and Wippich 1995: fig. 1). The lack of ribs on the body chamber inmacroconchs from the Kazimierz Dolny area versus their presence in material from Cotentin was the main criterion used by Błaszkiewicz (1980) to distinguish H. c. crassus from the nominal subspe− cies. This view is accepted herein, and all samples of H. constrictus in which smooth adult macroconchs predominate, are assigned to H. c. crassus. The subspecies is interpreted as part of the H. constrictus lineage intermediate between the earlier H. c. lvivensis subsp. nov. and the later H. c. johnjagti subsp. nov. (Fig. 8). The gradual character of the transition from H. c. crassus to H. c. johnjagti subsp. nov. is documented by the occasional occurrence of the ribbedmorphotype in pop− ulations of H. c. crassus from the upper part of the Kazimierz Opoka, as exposed at Nasiłów and Bochotnica (e.g., Machal− ski 1996: fig. 2E, 2005: fig. 10A, B), and in populations from the white chalk succession at Sigerslev quarry (Stevns Klint). This is further substantiated by smooth specimens in a sample of H. constrictus johnjagti subsp. nov. from the Grey Chalk of Stevns Klint (Machalski 2005; see also below) and by the presence of “transitional” specimens with weak ribbing, both from the Kazimierz Dolny area and from the “Dania” quarry (Machalski 2005). As pointed out by Machalski (2005), the name crassus was informally used by Birkelund (1979, 1993), Kennedy (1986, 1987), Jagt (1995), Machalski (1996) and other au− thors to denote specimens of Hoploscaphites constrictus with the body chamber entirely covered by ribs and with ventrolateral tuberculation extending to the aperture. Thus, the “crassus” concept of these authors was different from that of Błaszkiewicz (1980); his views are accepted in the present paper. In addition to the Middle Vistula River valley sections, the best material of H. c. crassus comes from a sample from the opoka layer between the belemnite and baculite marls at Rejowiec. This comprises 73 adult macro− and 15 adult microconchs, with the former ranging between 28 and 49 mm, the latter between 22 and 33 mm in maximum diameter, respectively (Fig. 9B). All macroconchs of this sample repre− sent the smooth morphotype (e.g., Fig. 10D), and do not reach the size of the largest specimens from the Kazimierz Dolny and Nasiłów samples (Fig. 10B–D). This difference may be ascribed to some ecophenotypic factors as proposed by Landman et al. (2003) for a similar size variation amongst samples of the North American scaphitid Hoploscaphites nicolletii (Morton, 1842). The remaining samples studied (see synonymy and mate− rial) are assigned to the subspecies based on population crite− ria, i.e., the predominance of macroconchs with smooth flanks of the shaft. These include samples from the opoka successions at Klimusin, Mętów, and Prawiedniki in the Lublin area and from the white chalk succession as exposed at Sigerslev quarry (Stevns Klint), described in more detail by Machalski (2005). Only specimen DPM NANU PZ−K− N9282 (Fig. 10C) from the “Lvovskaja svita” at Lviv was as− signed to this subspecies on a purely typological basis be− cause provenance data are lacking (see above). Alterna− tively, DPM NANU PZ−K−N9282 might represent an end− member of the population of H. c. lvivensis subsp. nov. Material from the upper, but not uppermost, Maastrich− tian of the Bay of Biscay region, referred to as Hoplo− scaphites constrictus (J. Sowerby, 1817) by Ward and Ken− nedy (1993: figs. 43.1, 43.2, 45.3) is too poorly preserved for subspecific assignment. The same concerns approximately coeval material of the species reported from Bjala (Bulgaria), by Ivanov and Stoykova (1994: pl. 3A, B). Adult microconchs of the subspecies show a remarkable, size−dependent variability in the ornament on the body cham− ber (Fig. 11). It seems as if the ornament was most complete in the largest individuals, encompassing a long interval of rather diffuse ribbing followed adaperturally by an interval of densely spaced ribs (Fig. 11F). The interval with diffuse ribbing seems to decrease in length in smaller−sized individuals (Fig. 11E), and is entirely lacking in the smallest specimens, where the adapertural stage of dense ribbing contacts directly with the regular ribbing typical of the phragmocone (Fig. 11D). The ob− served variation may be related to differential onset of matura− tion in specimens, with small individuals reaching adulthood earlier than large ones. This would concern only microconchs, interpreted as males (Makowski 1962). The macroconchs, thought to be females, do not show such size−dependent vari− ability in ornament (Fig. 11A–C). Such an explanation, if cor− rect,would lead to a rather unexpected merging of the hypothe− sis of sexual dimorphism in ammonites (Makowski 1962; Callomon 1963) with the developmental polymorphism theory as advocated by Matyja (1986). Unfortunately, neither whorls nor septa can be counted in the specimens studied, for preser− vational reasons. STRATIGRAPHIC AND GEOGRAPHIC RANGE.- Upper, but not upper− most,Maastrichtian of Poland (Spyridoceramus tegulatus– Be− lemnitella junior Zone and Belemnella kazimiroviensis Zone), Denmark (Argyrotheca stevensis–Magas chitoniformis Zone, Ruegenella humboldtii–Argyrotheca stevensis Zone and the equivalent Belemnella kazimiroviensis Zone) and the Ukraine (Spyridoceramus tegulatus–Belemnitella junior Zone). |
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| Systematics: | ||||||
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51 Familia Scaphitidae Genus Hoploscaphites Species Hoploscaphites constrictus Species Hoploscaphites schmidi Subspecies Hoploscaphites constrictus crassus Subspecies Hoploscaphites constrictus lvivensis |
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| Synonym list: | ||||||
| Machalski (2005): 1911 Scaphites constrictus var. crassus Lopuski. - Lopuski : p.115 pl.2, fig. 4-6;
pl. 3, fig. 1, 2 [var. nov.]
1911 Scaphites constrictus d'Orbigny. - Lopuski : p.113 pl. 2; fig. 3, 4
1911 Scaphites sp. . - Lopuski : p. 117, 118 pl. 3; fig. 3-6
p 1911 Hoploscaphites constrictus vulgaris Nowak. - Nowak : p.583 pl. 33; fig. 8-10 [non pl. 33; fig. 11, 12 =
Hoploscaphites tenuistriatus]
1962 Scaphites constrictus d'Orbigny. - Makowski : p.31 text-pl. 4; fig. 3
1962 Scaphites constrictus var. niedzwiedzkii Uhlig. - Makowski : text-pl. 4; fig. 4
1980 Hoploscaphites constrictus crassus Lopuski. - BLASZKIEWICZ : p.37 pl. 18; fig. 1-3, 11-14
p 1993 Hoploscaphites constrictus Sowerby. - BIRKELUND : p.57 pl. 14, fig. 5;
pl. 17, fig. 10 [non pl. 14, fig. 1-4, 6, 7, 12; pl. 15, fig. 1-14;
pl. 16, fig. 6-16; pl. 17, fig. 5-9, 11-23 =
Hoploscaphites constrictus johnjagti subsp. nov.]
1996 Hoploscaphites constrictus Sowerby. - Radwanski : pl. 2:1, 2; fig. 1-11
1996 Hoploscaphites constrictus Sowerby. - Machalski : fig. 2A-G, 3A-C
2005 Hoploscaphites constrictus crassus Lopuski. - Machalski : p. 665, 668 fig. 7A, D, G, H, 10
2005 Hoploscaphites ssp. constrictus A Machalski. - Machalski : fig. 8A-E, 10A, B, 11A-D
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| Was used in synonym list of: | ||||||
| Stratigraphy - relative ages: | ||||||
| upper Maastrichtian: Machalski (2005) |
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| References: | ||||||
Nowak,K.. (1911): Lopuski,C.. (1911): Makowski,H.. (1962): BLASZKIEWICZ,A.. (1980): BIRKELUND,T.. (1993): Radwanski,A.. (1996): Machalski,M.. (1996): Machalski,M.. (2005): |
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| Anonymous: Unedited TaxonConcept data | ||||||
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