by Thomas Brühwiler and Hugo Bucher
Systematic descriptions mainly follow the classification established by Tozer (1981, 1994) and refined by Brayard and Bucher (2008), Brühwiler et al. (2010b) and Brühwiler and Bucher (2011a, b).
Abbreviations: non = material not forming part of the current species; v. = video or vidimus (from Latin, means that the material was seen in person by the authors); ? = questionable; PIMUZ, Paläontologisches Institut und Museum der Universität Zürich, Switzerland.
Class Cephalopoda Cuvier 1797
Subclass Ammonoidea Zittel 1884
Order Ceratitida Hyatt 1884
Superfamily Xenodiscaceae Frech 1902
Family Kashmiritidae Spath 1934
Genus
Kashmirites
D
iener
1913
Type species. Celtites armatus Waagen1895.
Kashmirites weisserti
sp. nov.
Plate 1, Figs. 1–10
Derivation of name. Named for Helmut Weissert (Zürich).
Holotype. Specimen PIMUZ 28552 (Pl. 1, fig. 1a–d).
Type locality. Nammal, Salt Range, Pakistan.
Type horizon. Sample Nam802, top of CS, Euflemingites cirratus beds.
Diagnosis. Kashmirites with high intraspecific variation regarding strength of ornamentation and whorl section; ornamentation consists of simple ribs that become weak on the ventral shoulders but cross the venter.
Occurrence. Abundant in sample Nam802.
Description. Very evolute shell with slightly flattened flanks exhibiting large intraspecific variation. Whorls vary from slightly compressed to depressed. Venter, broad and subtabulate with rounded shoulders. Umbilicus, wide with inclined to almost vertical wall and rounded shoulders. Ornamentation consists of fine to very strong, straight ribs that become weak on the ventral shoulders but cross the venter. Ribs become fine and dense on the outer whorls. Suture line is with weakly indented lobes and broad saddles; third lateral saddle crosses the umbilical suture.
Measurements. See Fig. 6; Appendix.
Discussion. Variants of this species with weak ornamentation exhibit a slightly more involute coiling as well as a more compressed whorl section than variants with strong ornamentation. This type of covariation is well known as Buckman’s first law of covariation (Westermann 1966; Hammer and Bucher 2005). Kashmirites weisserti resembles K. baidi Brühwiler and Bucher (2011b) that has recently been reported from Oman as well as from the Clypeoceras superbum beds of the Salt Range (Brühwiler et al. 2011a, b). However, that species differs by its ornamentation consisting of simple or paired strong, radial ribs. The type species, K. armatus (Waagen1895), clearly differs from K. weisserti by its suture line with relatively strongly indented lobes and slightly phylloid saddles as well by its less sharp ribs and its smooth venter.
Family Flemingitidae Hyatt 1900
Genus
Flemingites
W
aagen
1895
Type species. Ceratites flemingianus De Koninck 1863.
Flemingites flemingianus(De Koninck 1863)
Plate 3, fig. 5.
1863 Ceratites flemingianus
de Koninck, p. 10, pl. 7, fig. 1.
1895 Flemingites flemingianus Waagen, p. 199, pl. 12, fig. 1, pl. 13, fig. 1, pl. 14, fig. 1.
1895 Flemingites compressus Waagen, p. 202, pl. 15, fig. 1, pl. 16, fig. 1a–c.
1895 Flemingites trilobatus Waagen, p. 193, pl. 26, fig. 2a–b.
1933 Flemingites compressus Waagen; Collignon, p. 170, pl. 3, fig. 1, 1a.
1933 Flemingites flemingi madagascariensis Collignon, p. 173, pl. 5, fig. 1.
1933 Flemingites griesbachi Krafft
and Diener; Collignon, p. 176, pl. 4, fig. 3, 3a; pl. 5, fig. 4; pl. 6, fig. 1–2.
v non 2008 Flemingites flemingianus Waagen; Brayard and Bucher, p. 43, pl. 17, figs. 1–5; text-fig. 38.
v acc. Flemingites flemingianus Waagen; Brühwiler et al.
Occurrence. Three specimens from sample Nam802. This species has previously been reported from a horizon ca. 2 m below this sample at Nammal as well as from several other localities in the Salt Range (Waagen 1895, Brühwiler et al. 2011b; Fig. 2).
Description. Large, evolute shell with elliptical whorl section. Venter, rounded with indistinct shoulders. Umbilicus, wide with rounded wall. Flanks, ornamented with strigation and radial or rursiradiate ribs. Ribs are strongest on inner whorls, but become finer in the outer whorls. Suture line, not preserved.
Discussion. See Brühwiler et al. (2011b) for a recent discussion of this species based on new and well-preserved material from the Salt Range.
Genus
Euflemingites
S
path
1934
Type species. Flemingites guyerdetiformis Welter 1922.
Euflemingites cirratus
(W
hite
1880)
Plate 2, figs. 1–4
1880 Arcestes cirratus White, p. 116.
1932 Flemingites cirratus White - Smith, p. 53, pl. 20, fig. 1; pl. 26, figs. 1–12.
? 1947 Flemingites prynadai Kiparisova, p. 135, pl. 29, fig. 1; pl. 30, fig. 1; text-fig. 18.
? 1954 Flemingites prynadai Kiparisova - Kiparisova and Krishtofovich, p. 18, pl. 8, fig. 1.
1959 Euflemingites tsotengensis Chao, p. 209, pl. 5, figs. 1–2.
? 1961 Flemingites prynadai Kiparisova - Kiparisova, p. 76, pl. 15, fig. 1a–b, text-fig. 36.
1967 Euflemingites cf. cirratus White - Tozer, p. 19, 50, 74.
1970 Euflemingites cf. cirratus White - Korchinskaya, p. 82, pl. 2, fig. 1.
1979 Euflemingites cirratus White - Nichols and Silberling, p. B3, pl. 3, figs. 1–3.
1982 Euflemingites cf. cirratus White - Korchinskaya, pl. 5, fig. 1; pl. 6, fig. 3.
1994 Euflemingites cirratus White - Tozer, p. 72, pl. 22, fig. 13; pl. 23, figs. 4–5, 21b.
? 2004 Euflemingites cf. tsotengensis Chao - Tong et al., p. 200, pl. 2, figs. 13–15.
? 2007 Euflemingites sp. - Krystyn et al., pl. 4, figs. 1–2.
? 2009 Eulemingites prynadai Kiparisova - Shigeta and Zakharov, p. 100, figs. 87–90.
Occurrence. Four specimens from sample Nam802.
Description. Moderately evolute, compressed shell. Flanks, flat and parallel. Venter, rounded and slightly flattened with rounded shoulders. Umbilicus, relatively deep with vertical wall. Ornamentation consists of very coarse strigation on the venter, flanks and umbilical wall. Suture line, ceratitic with long, slightly phylloid saddles and broad, strongly indented lobes.
Measurements. See Appendix.
Discussion. Our new material from the Salt Range is clearly conspecific with the well preserved specimens of Euflemingites cirratus from western USA described by Smith (1932). E.tsotengensis Chao 1959 from South China does not exhibit any differences from this species and is here considered as a synonym. The specimens described as E. cf. tsotengensis Chao 1959 by Tong (2004) are too poorly preserved for identification at the species level. E. prynadai (Kiparisova1947) from Primorye is very similar to E. cirratus and essentially differs by its larger size. The type species E. guyerdeti (Diener 1897), of which only a single small specimen from Spiti is known, differs slightly from E. cirratus by its inclined umbilical wall. The specimens from the Flemingites beds of Spiti described as Euflemingites sp. by Krystyn et al. (2007) may be conspecific with either E. guyerdeti or E. cirratus, but they are too poorly preserved for identification at the species level. E. guyerdetiformis (Welter 1922) from Timor differs from E. cirratus by its denser spiral ornamentation. E. cirratus has also been reported from the Chulitna Terrane in Alaska, from NE British Columbia and from Spitsbergen (Tozer 1967, 1994; Korchinskaya 1970, 1982; Nichols and Silberling 1979). The boreal species E. romunduri Tozer 1961 as well as E. aryomensis Smyshlyaeva2010 from Primorye differ from E. cirratus by their thicker whorl section.
Family Arctoceratidae Arthaber 1911
Genus
Arctoceras
H
yatt
1900
Type species. Ceratites polaris Mojsisovics 1886.
Arctoceras schalteggeri
sp. nov.
Plate 4, figs. 1–2
Derivation of name. Named for Urs Schaltegger (Geneva).
Holotype. Specimen PIMUZ 28563 (Pl. 4, Fig. 1a–e).
Type locality. Nammal, Salt Range, Pakistan.
Type horizon. Sample Nam802, top of CS, Euflemingites cirratus beds.
Diagnosis. Compressed Arctoceras without true tubercles but with weak ribs thickening at umbilical margin; and with subtabulate venter.
Occurrence. Relatively abundant in sample Nam802.
Description. Moderately evolute, compressed shell. Flanks nearly flat with a slight depression near the umbilical margin, only slightly converging. Venter subtabulate with rounded shoulders. Umbilicus deep with vertical wall and marked but rounded shoulders. Ornamentation consists of slightly biconcave ribs that are most prominent near the umbilical margin, where they almost develop tubercles. Suture line ceratitic, third lateral saddle low and broad, auxiliary series short.
Measurements. See Appendix.
Discussion. This species is very similar to the slightly younger Arctoceras meridionale Guex1978 from the lower part of the UCL of the Nammal section, which differs by its less platycone flanks, its more evolute coiling and its stronger, but very similar, ornamentation. That species is actually very close to Brayardites Brühwiler et al. (2010b) and may represent a transition form between the genera Arctoceras and Brayardites.
Typical Arctoceras from the paleoarctic realm differ from A. schalteggeri by their rounded venter and their more distinct tuberculation (e.g., Kummel 1961). A. tuberculatum (Smith
1932) from the Western USA and A. subhydaspis (Kiparisova
1961) are similar but differ by their more distinct tuberculation. A. strigatus Brayard and Bucher
2008 is very similar, but differs by its strigation, absence of umbilical tuberculation and a suture line with longer and slenderer saddles.
Family Galfettitidae Brühwiler and Bucher (2011b)
Genus
Vercherites
B
rühwiler
,W
are
, B
ucher
, K
rystyn
& G
oudemand
(2010b)
Type species. Koninckites vercherei Waagen, 1895.
Vercherites wyleri
sp. nov.
Plate 3, figs. 1–4
Derivation of name. Named for Daniel Wyler (Zürich).
Holotype. Specimen PIMUZ 28558 (Pl. 3, fig. 1a–d).
Type locality. Nammal, Salt Range, Pakistan.
Type horizon. Sample Nam802, top of CS, Euflemingites cirratus beds.
Diagnosis. Vercherites with tabulate venter on the inner whorls, which becomes rounded on the outer whorls; flanks with weak plications; suture line with a relatively short auxiliary series.
Occurrence. Rather abundant in sample Nam802.
Description. Moderately evolute, compressed shell. Flanks, convex and convergent. Venter, narrow and tabulate with slightly rounded shoulders on the inner whorls, becoming broader and rounded on the outer whorls. Umbilicus with vertical wall and marked but rounded shoulders. Surface, smooth except for slightly biconcave growth lines and weak plications that are most prominent at the mid-flanks. Suture line, simple and ceratitic with rather weakly indented lobes and a short auxiliary series.
Measurements. See Appendix.
Discussion. The older Vercheritesvercheri (Waagen1895) from the CM and V. pulchrum (Waagen1895) from the CS are similar but differ by their narrower and more tabulate venters (Brühwiler et al. 2011b). Paranorites ambiensis (Waagen1895) differs by its distinct suture line with a narrow and phylloid first lateral saddle. V. wyleri sp. nov. differs from Arctoceras schalteggeri sp. nov. described above by its different type of ornamentation and its gently curved flanks. Lepiskites Dagys & Ermakova 1990 differs by its ribbed inner whorls and its suture line with a long auxiliary series.
Family Melagathiceratidae Tozer 1971
Genus
Juvenites
S
mith
1927
Type species. Juvenites kraffti Smith 1927.
Juvenites
sp. indet.
Plate 4, fig. 3a–e
Occurrence. A single, very small specimen from sample Nam802.
Description. Moderately involute, subglobose shell. Venter, broadly rounded. Umbilicus, deep with vertical wall and well-rounded shoulders. Ornamentation consists of strong constrictions. Suture line, ceratitic with only two lateral saddles and weakly indented lobes.
Measurements. See Appendix.
Discussion. Our single specimen is possibly conspecific with the slightly younger Juvenites sp. indet. from the Brayardites compressus beds at Nammal (Brühwiler et al. 2011b), but its small size hinders a specific assignment.
Family incertae sedis
Genus
Kraffticeras
B
rühwiler
et
al
. (
2011a
)
Type species. Meekoceras pseudoplanulatum Krafft & Diener 1909.
Kraffticeras pseudoplanulatum
(K
rafft
& D
iener
1909)
Plate 4, fig. 5a–e
1909 Meekoceras pseudoplanulatum Kraft & Diener, p. 30, pl. 6, fig. 3a–c
acc. Kraffticeras pseudoplanulatum (Krafft & Diener 1909) - Brühwiler et al. (2011a).
Occurrence. Two small specimens from sample Nam802.
Description. Moderately involute, compressed shell. Flanks, nearly flat, only slightly convex and convergent. Venter, bicarinate. Umbilicus, deep with vertical wall and subangular shoulders. Surface, ornamented with weak plications that are most prominent near the umbilical margin. Suture line with long saddles and deep, apparently not indented lobes.
Measurements. See Appendix.
Discussion. This species has recently been described from the upper part of the Flemingites beds at Losar, Spiti (Brühwiler et al. 2011a). Thus, it is an important taxon for the correlation of the ammonoid succession from the Salt Range with that of Spiti.
Superfamily Sagecerataceae Hyatt 1884
Family Hedenstroemiidae Waagen
1895
Genus
Hedenstroemia
W
aagen
1895
Type species. Ceratites hedenstroemi Keyserling 1845.
?Hedenstroemia
sp. indet.
Plate 4, fig. 4a–c
Occurrence. A single fragmentary specimen from sample Nam802.
Description. Very involute, compressed shell. Flanks, convergent. Venter, narrow, probably slightly bicarinate, but poorly preserved. Umbilicus, small and deep with vertical wall and subangular shoulders. Surface, smooth except for fine, distinctly biconcave growth lines. Suture line not preserved.
Discussion. Our single specimen is identical to the inner whorls of Hedenstroemiaevoluta Spath 1934 from the Clypeoceras superbum beds of the Salt Range (Brühwiler et al. 2011b). However, since the suture line is not preserved in this specimen, the assignment to Hedenstroemia is only provisional.