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TABLE 1. Overview of species used in the analysis. Assignment to suborders is here based on previous taxonomic concepts, which may differ from the results obtained here. For more details, see supplementary material.

Species (Sub-) order Age Main sources
Atractites alpinus von Gümbel, 1861 Aulacoceratida Early Jurassic  Jeletzky, 1966; Mariotti et al., 2021
Aulacoceras sulcatum  von Hauer, 1860 Aulacoceratida Late Triassic Jeletzky, 1966; Mariotti et al., 2021
Acrocoelites oxyconus (Hehl in von Zieten, 1831) Belemnitina Early Jurassic Schlegelmilch, 1998
Aulacoteuthis ernsti Mutterlose and Baraboshkin, 2003 Belemnitina Early Cretaceous Mutterlose, 1983; Mutterlose and Baraboshkin, 2003
Cylindroteuthis puzosiana (d’Orbigny, 1842) Belemnitina Late Jurassic Schlegelmilch, 1998
Megateuthis gigantea (von Schlotheim, 1820) Belemnitina Middle Jurassic Schlegelmilch, 1998, own data
Oxyteuthis brunsvicensis (Strombeck, 1861) Belemnitina Early Cretaceous Stolley, 1911b; Mutterlose, 1983
Passaloteuthis bisulcata (Blainville, 1827) Belemnitina Early Jurassic Schlegelmilch, 1998
Schwegleria feifeli (Schwegler, 1939) Belemnitina Early Jurassic Schlegelmilch, 1998
Belemnitella mucronata (von Schlotheim, 1813) Belemnopseina Late Cretaceous Christensen, 1997
Belemnitella propinqua (Moberg, 1885) Belemnopseina Late Cretaceous Christensen, 1971
Belemnopsis apiciconus (Blainville, 1827) Belemnopseina Middle Jurassic Schlegelmilch, 1998
Calabribelus pallinii  Weis et al., 2012 Belemnopseina Middle Jurassic Weis et al., 2012
Dicoelites dicoelus (Boehm, 1906) Belemnopseina Late Jurassic Stevens, 1964
Dimitobelus diptychus (McCoy, 1867) Belemnopseina Early Cretaceous Whitehouse, 1924; Williamson, 2006
Duvalia grasiana (Duval-Jouve, 1841) Belemnopseina Early Cretaceous Stolley, 1911a; Stoyanova-Vergilova, 1970; Combémorel, 1973; own ata
Gonioteuthis quadrata (Blainville, 1827) Belemnopseina Late Cretaceous Ernst, 1964; Christensen 1997
Hibolithes semisulcatus (zu Münster, 1830) Belemnopseina Late Jurassic Schlegelmilch, 1998; own data
Holcobelus munieri (Eudes-Deslongchampes, 1878) Belemnopseina Middle Jurassic Jeletzky, 1966; Weis et al., 2012
Lissajousibelus harleyi (Mayer, 1866) Belemnopseina Early Jurassic Weis et al., 2015b
Mesohibolites minaret (Raspail, 1829) Belemnopseina Early Cretaceous Stoyanova-Vergilova,1970
Neohibolites ewaldi (Strombeck, 1861) Belemnopseina Early Cretaceous Stolley, 1911a; own data
Neohibolites minimus (Miller, 1826) Belemnopseina Early Cretaceous Stolley, 1911a; Spaeth, 1971; Stevens et al., 2017
Palaeobelemnopsis sinensis (Chen, 1982) Aulacoceratida incertae sedis Late Permian Mariotti et al., 2021
Praeactinocamax plenus (Blainville, 1827) Belemnopseina Late Cretaceous Christensen, 1997
Sinobelemnites cornutus Zhu and Bian 1984 Belemnopseina Late Triassic Zhu and Bian 1984
Tohokubelus takaizumii Niko and Ehiro, 2022 Belemnopseina Early Triassic Niko and Ehiro 2022

 

 

TABLE 2. List of included characters. Representation of hierarchical character state dependencies follows Pohle et al. (2022), i.e., subordinate characters are scored as “inapplicable” (-), if higher level character is scored as state 0. See also Figure 1 for examples.

# Character description Character states
1 Two dorsolateral apical furrows absent (0), present (1)
2 Apical ventral furrow absent (0), present (1)
3 Two ventrolateral apical furrows absent (0), present (1)
4 Ventral alveolar furrow(s) absent (0), present (1)
4.1 Ventral alveolar furrow type intermediate (0), reaching alveolus (1)
4.1.1 Ventral alveolar furrow number one (0), two (1)
4.1.2 Ventral furrow reaches from alveolus towards apical region no (0), yes (1)
4.2 Ventral splitting surface (slit, slitfield, “Schlitzfeld”) absent (0), present (1)
4.2.1 Ventral splitting surface type rudimentary (0), full (1)
5 Dorsal alveolar furrow absent (0), present (1)
5.1 Dorsal splitting surface (slit, slitfield, “Schlitzfeld”) absent (0), present (1)
6 Doppellinien absent (0), present (1)
6.1 Doppellinien type parallel (0), diverge towards alveolus (1), multiple (2)
7 Proper rostrum mineral aragonite (0), calcite (1)
8 Pseudoalveolus, secondary erosion of alveolus forming a concave or conical structure absent (0), present (1)
8.1 Pseudoalveolus type convex (0), concave (1)
9 Alveolus angle low, < 11° (0), high > 11°
10 Alveolus type central (0), ventrally displaced (1)
11 Rostrum surface ribbed with longitudinal ridges (0), smooth (1)
12 Two dorsolateral longitudinal depressions absent (0), present (1)
13 “Vascular” imprints absent (0), present (1)
14 Apex type acute (0), obtuse (1)
15 Epirostrum absent (0), present (1)
16 Cameral deposits absent (0), present (1)
17 Juvenile rostrum type conirostrid (0), clavirostrid (1)
18 Rostrum hastate in dorsal view no (0), yes (1)
19 Deep lateral furrows absent (0), present (1)
20 Cross-section at position of protoconch +/- circular (0), compressed (1)
21 Ventral tongue-like extension absent (0), present (1)

 

 

 

TABLE 3. Results of the parameter estimates. Abbreviations: stdev = standard deviation, HPD int = highest posterior density interval, SA = sampled ancestors.

Parameter mean median stdev 95% HPD int
Clock rate 0.0052 0.0051 0.0013 0.0030 - 0.0075
Diversification 0.012 0.011 0.0069 0.00084 - 0.025
Turnover 0.918 0.926 0.049 0.821 - 0.992
Sampling 0.075 0.073 0.039 0.015 - 0.147
Origin 270.6 269.3 8.63 256.5 - 288.1
SA count 4.71 5 1.98 1 - 8

 

 

 

TABLE 4. Proposed revised systematic scheme for the Belemnitida based on the Bayesian tip-dated analyses. For families, we follow previous systematic concepts (see also Table 1), although some were recovered as paraphyletic, indicated with (*). Note that only families that were included in the analyses are listed here. Aulacoceratids are omitted here because they were not a central focus of the study, providing only limited insights for their classification.

Class Cephalopoda Cuvier, 1797
  Subclass Coleoidea Bather, 1888
    Order Belemnitida Zittel, 1895
          Family Sinobelemnitidae (*) Bian and Zhu, 1984
      Suborder Belemnitina Zittel, 1895
          Family Passaloteuthidae (*) Naef, 1922
          Family Megateuthidae Sachs and Nalnjaeva, 1967
      Suborder Belemnopseina Jeletzky, 1965
          Family Holcobelidae Gustomesov, 1977
          Family Dicoelitidae Sachs and Nalnjaeva, 1967
          Family Duvaliidae Pavlov, 1914
          Family Cylindroteuthidae Stolley, 1919
          Family Oxyteuthidae Stolley, 1919
        Pseudoalveolata nov. [unranked]
          Family Belemnopseidae (*) Naef, 1922
          Family Dimitobelidae Whitehouse, 1924
          Family Belemnitellidae Pavlow, 1914