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 |